Dominik Schüssler

Marina B Blanco

Jordi Salmona

Jelmer Poelstra

Jean B Andriambeloson

Alex Miller

Blanchard Randrianambinina

David Rasolofoson

Jasmin Mantilla-Contreras

Lounès Chikhi

Dominik Schüßler

W Rasolofoson

Anne D Yoder

Ute Radespiel
email: ute.radespiel@tiho-hannover.de

Ecology and morphology of mouse lemurs ( Microcebus spp.) in a hotspot of microendemism in

Keywords: evolution, cryptic species, phenotype, habitat use, sympatry evolution, cryptic species, phenotype, habitat use, sympatry, 80 conservation 81

de niveau recherche, publiés ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

116

Recent studies indicate that some regions appear to be hotspots of 117 microendemism. One of these is located in northeastern Madagascar where M. 118 lehilahytsara (Kappeler et al., 2005), M. mittermeieri and M. simmonsi (Louis et 119 al., 2006) are known to occur. Radespiel et al. (2008) surveyed the forests of 120 the Makira region (Anjiahely, Figure 1) and found evidence for three divergent 121 lineages occurring in sympatry, a phenomenon previously undocumented for 122 mouse lemurs. One of these was identified as M. mittermeieri, while the second 123 was newly described as M. macarthurii. The third lineage, named M. sp. #3, 124 was hypothesized to be a new species based on mitochondrial sequence data 125 but could not be formally described given that only a single individual was 126 found.

127

We conducted additional sampling in northeastern Madagascar to fill the gap 128 between the known distribution of M. simmonsi (Zahamena NP, Betampona 129 SNR, Tampolo; Louis et al., 2006) and the sympatric species pair M. 130 macarthurii and M. mittermeieri at Anjiahely (Radespiel et al., 2008; Figure 1).

131

The presence of the M. sp. #3 lineage was indeed confirmed but only for three 2). Instead, these 144 two latter species exhibited a single isolation-by-distance pattern and high 145 levels of interspecific gene flow, suggesting that separate species status may 146 not be justified (Figure 2; discussed in detail in Poelstra et al., 2020). For the 147 purpose of this study, however, we will still treat these two lineages as separate 148 taxa in order to be able to test their distinctiveness in other domains. rainforests as primary vegetation (Kottek, Grieser, Beck, Rudolf & Rubel, 2006).

167 Forest cover has been steadily declining for decades, with lowland rainforests 168 being particularly prone to deforestation (Schüßler,169 Stadtmann, [START_REF] Schüßler | Identification of crucial stepping stone habitats for Page 44 of 66[END_REF]Vieilledent et al., 2018). By 2018, 170 about half of the remaining forested areas were under protection by 171 governmental institutions or non-governmental organizations (Schüßler et al., 172 2020). The study region is subdivided by more than seven large rivers that flow 173 from the highlands of the central plateau (west of the study region; Figure 1) the vicinity of Mananara-Nord NP, as well as in the Ambodiriana community 186 protected area (Miller et al., 2018). All study sites comprise habitats ranging 187 from undisturbed near-primary rainforest to heavily degraded secondary shrub-, 188 grass-and fernlands (Radespiel et al., 2008;Miller et al., 2018;[START_REF] Schüßler | Identification of crucial stepping stone habitats for Page 44 of 66[END_REF] S1).

229

Mouse lemurs are seasonal breeders and can already reproduce during their 230 first year of life (Evasoa et al., 2018;Kraus, Eberle & Kappeler, 2008; 231 [START_REF] Schmelting | Experience-dependent recapture rates and reproductive success in 1002 male grey mouse lemurs (Microcebus murinus)[END_REF] S3) and Tukey post-hoc tests revealed many pairwise differences 314 (Figure 3; Table 3; Figure S1). M. sp. #3 can be statistically differentiated from 315 its closest relative, M. macarthurii, by 5 out of 13 parameters. M. macarthurii 316 has smaller body size and longer tail length, and subtle differences were found 317 in head-associated parameters (i.e., ear width, head length and width). By 318 comparison, M. lehilahytsara (from Mantadia NP) and M. mittermeieri (from 319 Anjiahely) differed in 7 out of 13 variables. Major differences were found in 320 snout and tail length, while other differences were more subtle but statistically 321 significant (Figure 3; Figure S1; Table 3). Both M. lehilahytsara and M. 322 mittermeieri were significantly smaller than M. sp. #3 and M. macarthurii, which 323 was mainly reflected in the parameters body mass and length, tail length, lower 324 leg, hind foot and third toe length (Figure 3, Figure S1; Table 3). M. simmonsi [START_REF] Schneider | Signals of recent 1005 spatial expansions in the grey mouse lemur (Microcebus murinus)[END_REF]. Here, we confirm a new case 402 with M. sp. #3 being found in sympatry with M. lehilahytsara in Ambavala

403

(Figure 1).

404

Ecologically, two sister species, M. macarthurii and M. sp. #3, appear to be 405 restricted to lowland forests, whereas M. mittermeieri and M. lehilahytsara are 406 present in lowland as well as montane forests ( Radespiel, 2016). However, the recent expansion of M. murinus seems to have 415 been a unique event that has no clear equivalent in eastern Madagascar.

416

M. simmonsi was previously reported from Zahamena NP, Betampona SNR 417 and Tampolo (Hotaling et al., 2016;Louis et al., 2006). We can now confirm its 418 occurrence 75 km and four inter-river systems (IRSs) further north (Figure 1 & 419 2) which expands its EOO by almost fivefold. The northern range limit for M. 420 simmonsi appears to be the Anove River, which separates it from M. sp. #3.

421

These two species have (so far) only been found in allopatry (despite intensive 422 sampling north of the river), and we consider two alternative hypotheses 423 responsible for this pattern: (1) competitive exclusion at the geographic limits of 424 the respective species ranges (Beaudrot et al., 2013;Hardin, 1960) River, however, may have been precluded by the presence of the larger M. sp.

433

#3 that may have a higher competitive potential than the smaller M. simmonsi 434 (Table 3;Thorén, Linnenbrink & Radespiel, 2011). In the case of M. sp. #3, the 435 subpopulations on both sides of the large Mananara River (Figure 1) were 436 shown to belong to two separate population clusters evolving largely 437 independently from each other (Figure 2; Poelstra et al., 2020). This suggests 438 that the Mananara River poses a significant barrier to gene flow within this 439 species (Poelstra et al., 2020) Mouse lemurs are typically regarded as cryptic species exhibiting only subtle 467 interspecific morphological differences [START_REF] Zimmermann | Species concepts, diversity, and 1060 evolution in primates: Lessons to be learned from mouse lemurs[END_REF].

468

Although our measurements of 13 external body parameters generally confirm 469 their cryptic nature, some differences could be detected that can help to 470 distinguish different species.

471

All analyses confirmed a noticeable divide between the two larger taxa M. sp.

472

#3/M. macarthurii and the two smaller-bodied lineages M. lehilahytsara and M. length, third toe length and four head-associated parameters (Figure 3; Table 489 3). This comparison was, however, based on two populations that are over 400 490 km away from each other. Genomic analyses on samples obtained for M. 3). If considered as one species, these differences could indicate 502 morphological adaptations to different environmental conditions (highland 503 versus lowland rainforest) or a morphological gradient across its entire range.

504

These hypotheses, however, will require further testing.

505

M. simmonsi individuals collected at Ambodiriana fell in between the clusters of 506 larger and smaller-bodied species in all analyses. However, when comparing 507 these individuals to M. simmonsi sampled further south (Zahamena NP and 508 Betampona SNR; Louis et al., 2006), our individuals were smaller (9.2 vs. 8.9 509 cm body size), had lower body mass (65 vs. 52 g) and had shorter tails (14.2 510 vs. 13.1 cm). These differences are even more pronounced when comparing 511 our dataset to the holotype specimen for M. simmonsi (9.8 cm, 77 g and 14.9 August/September, while females were still anestrous and not lactating (Figure 537 6), and juvenile individuals were not found. This timing of reproduction is 538 comparable to that of M. lehilahytsara from other sites but interestingly not from 539 Ambavala, where this species is sympatric with M. sp. #3. In September, the 540 two adult male M. lehilahytsara from Ambavala did not have enlarged testes December. An adult male captured in early March showed evidence of tail 558 fattening. At Tsinjoarivo (Table S2), another high-altitude forest, M. lehilahytsara 559 females were observed gestating or lactating in late November, December and 560 early January (Blanco, 2010). From these sites there was also evidence of Microcebus jonahi is a large-bodied, reddish-brown and small-eared mouse 709 lemur (Figure 7). This species has short and dense fur. The head is rufous

  of the world's prime biodiversity hotspots and its endemic 84 group of primates, the lemurs (Primates; Lemuriformes), are flagships for 85 species conservation(Myers, Mittermeier, Mittermeier, Da Fonseca & Kent,

132

  Figure1), these analyses unambiguously supported four different lineages: M.

149

  Besides using molecular data, species delimitation under an integrative 150 taxonomic approach (e.g., Padial, Miralles, de la Riva & Vences, 2010) requires 151 incorporating morphological and ecological information. While phenotypic 152 differences between lineages could, for instance, indicate dietary preferences 153 (e.g., Viguier, 2004), environmental and reproductive data can help to 154 understand the role of habitat selection (Dammhahn & Kappeler, 2008; 155 Rakotondravony & Radespiel, 2009) and reproductive schedules (Evasoa et al. 156 2018) during speciation. 157 Here, we complement the molecular results presented by Poelstra et al. (2020) 158 by (1) providing morphological, ecological and distributional data for the M. sp. 159 #3 lineage in comparison to all other species from the same region and (2) by 160 formally describing this new species.

  characterized by a humid climate with abundant 165 precipitation (2,086 ± SD 530 mm SD; Fick & Hijmans, 2017) and tropical 166

174

  eastwards into the Indian Ocean. Large rivers have been considered potential 175 biogeographic boundaries for mouse lemurs (e.g.,Olivieri et al., 2007; Martin,

  captured using Sherman Live traps (H. B. Sherman Traps 191 ®) or by hand during nocturnal surveys (e.g., Radespiel et al., 2008). 192 Morphometric measurements were taken for each individual (see below) and 193 additional descriptors such as fur coloration were noted and photographed. Ear 194 biopsies (~2 mm²) were collected to provide DNA samples, and all animals were 195 released unharmed within 24 hours at their exact location of capture. 196 GPS coordinates and the altitude of capture locations were collected to 197 estimate the altitudinal range and Extent of Occurrence of mouse lemur species 198 included in this study. The latter measure follows the definition of the IUCN 199 (2012) in which the "shortest continuous imaginary boundary which can be 200 drawn to encompass all the known, inferred or projected sites of present 201 occurrence of a taxon" is used to derive the possible distribution of a certain were measured for 13 different morphometric variables 212 (ear length, ear width, head length, head width, snout length, inter-and intra-213 orbital distance, lower leg length, hind foot length, third toe length, tail length, 214 body length and body mass) following Hafen, Neveu, Rumpler, Wilden and 215 Zimmermann (1998) and Zimmermann et al. (1998). Mouse lemurs were 216 assigned to two age categories based on their body mass and reproductive 217 state: (1) adult in contrast to (2) young mouse lemurs (<1 year old) that had a 218 relatively low body mass (Table 1) and showed no visible nipples (females) and 219 undeveloped, or barely developed testes (males). Young mouse lemurs were 220 excluded from all morphometric comparisons and only values for adult 221 Microcebus spp. are used for further analyses. Very light young mouse lemurs 222 of about half of the adult body mass were termed juveniles. 223 To supplement the comparative dataset, we also included published 224 morphometric data from 42 M. lehilahytsara individuals from Mantadia NP 225 (holotype locality, Randrianambinina, 2001) and data from 22 M. mittermeieri 226 individuals that had previously been caught near Anjiahely (Radespiel et al., 227 2008). The morphometric dataset is provided in the supplementary material 228 (Table

325

  took an intermediate position in most comparisons. M. lehilahytsara from 326 Ambavala was not compared by ANOVA due to the small number of adults (N = 327 3). However, the individuals from this population showed remarkably different 328 measures compared to conspecifics from Mantadia NP or M. mittermeieri from 329 Anjiahely (i.e., body length, tail length;

  simmonsi were caught in Ambodiriana in June and July. All males (N = 379 5) had enlarged testes, while females (N = 3) were not reproductively active 380 (Figure6).

  distribution, morphology and reproductive state of five mouse 384 lemur lineages occurring in a complex spatial pattern across a small region in 385 northeastern Madagascar. Comparative phenotypic and ecological data was 386 previously sparse for four of these taxa, and this study therefore represents an 387 important step towards deepening our understanding of mouse lemur diversity 388 in this understudied hotspot of microendemism.389 390 Distribution of Microcebus spp. in northeastern Madagascar 391 Our study, in conjunction with the companion study by Poelstra et al. (2020), 392 confirmed the presence of five Microcebus spp. lineages, including two pairs of 393 closely related sister lineages, M. macarthurii / M. sp. #3 and M. mittermeieri / 394 M. lehilahytsara in northeastern Madagascar, with two cases of local sympatry. 395 Besides the case of M. macarthurii and M. mittermeieri at Anjiahely (Radespiel 396 et al., 2008), sympatry of two mouse lemur species is so far only known from 397 five cases from western Madagascar and one case from the northern part of the 398 island (Sgarlata et al., 2019). In the five western cases, geographically 399 restricted species co-exist with the widely distributed congener M. murinus 400 (Radespiel, 2016) which probably expanded northwards rather recently

  473mittermeieri (along PC1, Figure4,

491

  lehilahytsara at two intermediate locations (Riamalandy and Ambavala, Figure 492 1) and M. mittermeieri samples from Anjiahely, Anjanaharibe-Sud SR and 493 Marojejy NP revealed only moderate genomic differentiation along a geographic 494 gradient (Figure 2; Poelstra et al., 2020). It was concluded that these results do 495 not justify separate species status of these two taxa. Unfortunately, we only 496 caught three adult M. lehilahytsara at Ambavala making it difficult to assess 497 whether the individuals from this intermediate geographic location also took an 498 intermediate morphometric position. Measurements from these three , suggest that M. lehilahytsara from Ambavala was more 500 similar to M. mittermeieri from Anjiahely than to M. lehilahytsara from Mantadia 501 NP (Table

  512 cm, respectively; Louis et al. 2006). On the other hand, the holotype specimen 513 of M. boraha, occurring on Ile St. Marie (an island less than 20 km off the coast 514 of Ambodiriana, Figure 1) had about the same average body mass (56.5 g; 515 Hotaling et al., 2016) as the population in Ambodiriana. While these 516 comparisons should be interpreted with caution due to small sample sizes and 517 different collection details, they suggest that M. simmonsi from Ambodiriana and 518 M. boraha from Ile St. Marie may be more closely related than previously 519 thought. Genomic analyses revealed only slight differentiation between 520 southern M. simmonsi and the population at Ambodiriana (Poelstra et al., 2020). 521 However, future analyses should also include samples from M. boraha from Ile 522 St. Marie to clarify relationships among these populations and lineages.

  Microcebus spp. in northeastern Madagascar 525 Differences in reproductive schedules among mouse lemur species depend on 526 phylogenetic relatedness, although environmental parameters (i.e., rainfall and 527 temperature) likely play a role in fine tuning reproductive function (Evasoa et al., 528 2018). Although photoperiod is generally considered the main trigger of 529 reproductive physiological function in mouse lemurs and is a relatively good 530 predictor of food availability in seasonal habitats, there is substantial variation in 531 the timing and duration of reproduction within and across mouse lemur species, 532 as evident in our study populations. 533 Reproductive observations from this study and those from published sources 534 (Evasoa et al., 2018) suggest that M. sp. #3 may start their mating season in 535 October-November, because males showed developed testes by 536

541

  and we also captured juvenile M. lehilahytsara with less than half of the adult 542 body mass (19-23 g). A body mass of about 20 g was found around the time of 543 weaning in juvenile captive M. lehilahytsara of about seven weeks of age 544 (Radespiel, Zimmermann & Wittkowski, unpubl. data). Adding these seven 545 weeks (49 days) to about 57 days of gestation (for M. lehilahytsara, 546 Wrogemann & Zimmermann, 2001; for M. rufus, Blanco, 2008), this could point season of the M. lehilahytsara in Ambavala potentially lasting 548 until May. 549 These findings contrast strikingly with those from M. lehilahytsara inhabiting 550 montane and high elevation rainforests. For instance, at Mantadia NP (300 km 551 further south of Ambavala; Figure 1) all adult males had enlarged testes in 552 September, and some adult females were in estrus in November. Juveniles at 553 this site (20 g body mass) were only found in late March (Randrianambinina et 554 al., 2003; this study). Similar observations were made at Ambatovy, a montane 555 forest (near Mantadia NP) where MBB (unpubl. data) captured two juveniles in 556 late February (~2 months old), suggesting a distinct birth season in mid-late 557

561

  rebound polyestry, i.e., females undergo renewed estrus after the loss of 562 offspring or early abortions. The capture of juvenile mouse lemurs (2-3 months 563 old) in early February, however, suggested a main birth season at this site in 564 early-mid December. In sum, observations of tail fattening at Tsinjoarivo and 565 Ambatovy in early March suggest that, at least for a portion of the mouse lemur 566 population, reproductive season is over by this time of the year. 567 Ambavala, unlike Mantadia NP, Ambatovy and Tsinjoarivo, is a low elevation 568 rainforest site (250 m vs. >800 m a.s.l., respectively). Seasonal climatic 569 fluctuations are more pronounced in montane rainforests than in the lowland 570 rainforests of eastern Madagascar. For example, Mantadia NP has a lower Schüßler et al. page 25 571 mean annual temperature (19°C compared to 23°C), lower minimum 572 temperature (9°C compared to 15°C) and a higher annual temperature range 573 (19°C compared to 15°C; Fick & Hijmans, 2017) than Ambavala. Climatic 574 conditions have already been suggested as major determinants of reproductive 575 schedules for small-bodied mouse lemurs (Evasoa et al., 2018) with higher 576 mean ambient temperatures and smaller temperature fluctuations allowing a 577 less seasonal reproduction compared to the harsh conditions of the montane 578 rainforests (Evasoa et al., 2018; Randrianambinina et al., 2003). Taken 579 together, the findings of this study and previous work suggest a considerable 580 degree of intraspecific variation concerning reproductive timing in M. 581 lehilahytsara. 582 Another pair of sympatric mouse lemur species from lowland rainforests 583 (Anjiahely: about 350-400 m a.s.l.) were characterized with regard to 584 reproductive activities in the northern part of our study region: M. macarthurii 585 and M. mittermeieri. The enlarged testes of M. macarthurii males from October 586 onwards and the presence of pre-estrous females in November indicate that the 587 mating season in these forests probably begins at that time (Figure 6). 588 Furthermore, the absence of juvenile and young mouse lemurs during our 589 sampling period may indicate one short mating season probably limited to 590 November-December. 591 Sympatric M. mittermeieri females were also found in estrus in November and 592 the respective males had enlarged testes in September. Moreover, juvenile M. 593 mittermeieri of about 20 g (~2 months old) were also found in November and 594 December. In analogy to the case of M. lehilahytsara (see above), we therefore Schüßler et al. page 26 595 also predict some mating of M. mittermeieri to occur as early as in July -August 596 in Anjiahely. Reproductive observations of two M. mittermeieri females from 597 another low elevation forest (Marojejy NP east site, ~200 m a.s.l.; MBB, unpubl. 598 data) captured in mid-late March, also suggest an extended reproductive period 599 and evidence of polyestry in this species. One of the females showed vaginal 600 swelling (pre-estrus) and 3 pairs of developed nipples, suggesting she had 601 given birth earlier in the season and was getting ready to mate again. The 602 second female showed a vaginal opening (estrus) and well-developed nipples, 603 also suggesting she had given birth earlier in the season and could, if mating 604 were successful, have another birth at the end of May or early June. 605 Interestingly, reproductive observations of M. mittermeieri from a montane 606 rainforest at Anjanaharibe Sud SR (980 m a.s.l; MBB, unpubl. data), were more 607 similar to those of M. lehilahytsara from other montane/high elevation forests 608 than to conspecifics from nearby low elevation forests: for instance, one adult 609 female captured in early May showed visible nipples but no signs of recent 610 lactation and evidence of tail fattening, and one adult male showed regressed 611 testes and tail fattening during the same capture period. Five captured young 612 mouse lemurs (~2-3 months old) showed no signs of reproductive activity and 613 no evidence of tail fattening. That a portion of the mouse lemur population is 614 displaying tail fattening in early May is indicative of a shorter reproductive 615 season, likely over by March. 616 In conclusion, and if supported by future studies, the M. lehilahytsara/M. 617 mittermeieri lineage in the warmer lowland rainforests of eastern Madagascar 618 seem to have more flexible and extended reproductive schedules compared to Schüßler et al. page 27 619 populations living in the harsher montane humid forest sites or to species 620 inhabiting the western dry deciduous forests (see Evasoa et al., 2018). In this 621 respect, these mouse lemurs show similarities to M. mamiratra and M. 622 margotmarshae that live in the lowland humid forests of northwestern 623 Madagascar (Sambirano region) and that are also reproductively active in the 624 first half of the dry season (Evasoa et al. 2018). Reproductive timing of 625 sympatric non-sister species was different, since the two microendemic species 626 (M. macarthurii and M. sp. #3) did not show extended reproductive schedules, 627 while allopatric sister linages showed highly similar schedules in both cases.

  five different Microcebus lineages were demonstrated to occur in a 631 geographically restricted region of northeastern Madagascar. All of these taxa 632 inhabit a 130 km wide stretch of lowland rainforest making this region one of the 633 most species-rich areas so far identified for mouse lemurs. These lineages can 634 be distinguished genetically (Poelstra et al., 2020) and morphologically as 635 shown in this study. The genomic and phenotypic differentiation between M. 636 macarthurii and M. sp. #3 provides sufficient support for the recognition of M. 637 sp. #3 as a distinct species which we describe below (see Species Description). 638 The studied taxa were found in a variety of habitat types, ranging from nearly 639 undisturbed to selectively logged forest, from shrubby secondary re-growth 640 vegetation to areas dominated by perennial plants (i.e., Aframomum spp.; Miller 641 et al., 2018; Radespiel et al., 2008; Schüßler et al., 2018). However, species 642 differed most substantially in their altitudinal range and the inferred Extent of

  Schüßler et al. page 28 643 Occurrence. M. lehilahytsara and M. mittermeieri were found both in montane 644 and lowland forests, but M. sp. #3 and M. macarthurii occurred only in lowland 645 forests. Locally restricted so-called "lowland specialists" have been found in at 646 least three other cases along the Malagasy east coast, i.e., M. gerpi (Radespiel 647 et al., 2012), M. marohita (Rasoloarison et al., 2013) and M. jollyae (Louis et al., 648 2006). These taxa share a narrow altitudinal range and a small estimated 649 Extent of Occurrence. Alarmingly, lowland rainforest habitats have disappeared 650 from most of the east coast and our study region is no exception (Schüßler et 651 al., 2020; Vieilledent et al., 2018). Under these circumstances, population 652 declines are unavoidable, and ongoing anthropogenic land use change and 653 forest cover loss (Schüßler et al., 2020) will probably accelerate and increase 654 this threat to the newly described species. M. sp. #3 occurs in one National 655 Park, Mananara-Nord NP, which is already isolated from surrounding forests. 656 Anticipated environmental disruptions due to future climatic changes and the 657 need for species to flexibly adapt to altered environments may be severely 658 compromised, if conservation planning does not generate and maintain possible 659 dispersal pathways for migrating species (Brown & Yoder, 2015; Schüßler et al., 660 2020).

  710colored with a darker brownish area around the eyes which can slightly vary 711 among individuals. A distinct white stripe lies between the eyes ending at the 7C). The ears are of the same rufous color as the head. The 713 cheeks are lighter brownish and less rufous than the head becoming even 714 lighter and almost white towards the throat. The ventrum is white with slightly 715 yellowish nuances (Figure 7D) which can vary in appearance among 716 individuals. The dorsum is rather uniformly brown than reddish (Figure 7E). A 717 darker dorsal stripe can be either present or absent. The ventrum and dorsum 718 are separated by a significant change in coloration with only marginal transition. 719 The coloration of the limbs shows the same pattern with a brownish dorsal and 720 a white to slightly yellowish ventral side. The tail is densely furred and of the 721 same coloration as the dorsum. Hands and feet show only sparse but whitish-722 gray hair. The skin on the palmar and plantar surfaces of hands and feet is 723 brownish pink. Males and females do not show any sexual dimorphism. individuals were captured at altitudes between 42 and 356 m a.s.l. 727 Out of the 25 captured individuals, six were caught in near-primary forest (= low 728 degradation) with a rather continuous canopy and five were captured in highly 729 degraded forests with discontinuous canopy cover and strong regrowth of early 730 successional trees. The majority of individuals (N = 14) were captured in 2 -4 m 731 high stands of the perennial Madagascar cardamom (Aframomum 732 angustifolium), sometimes intermixed with trees along the forest edges. At 733 Antanambe (Figure 1), a total of 39 individuals were sighted of which 35 were 734 found in forest habitats of different degradation stages. No M. jonahi were 735 sighted in treeless secondary vegetation except for dense Aframomum

Figure 1 : 1110 Figure 2 :Figure 4 :Figure 5 :Figure 6 : 1142 Figure 7 :

 11110245611427 Figure legends:

  . Reproductive ,250 km²) and M. lehilahytsara (50,700 km²; Table 2). Combined 308 EOO for M. lehilahytsara and M. mittermeieri, if considered as a single species,

	Schüßler et al. page 14
	307 mittermeieri (9309 is estimated with 66,800 km².
	F F F
	o r o r o r
	P P P
	e e r e e r e e r
	R e v R e v R e v
	i e w i e w i e w

232

states were assessed using several morphological indicators (i.e., vaginal 233 morphology and testis size) frequently used in the literature (e.g.,

Blanco, 2008;

Table 3

 3

).

  Table 2). Thus, M. macarthurii 407 and M. sp. #3 both have geographically restricted distributions and possess a 408 limited Extend of Occurrence, suggesting that they are microendemic, while M. sp. #3/M. lehilahytsara and M. macarthurii/M.

	411	mittermeieri, respectively, may to some extent be similar to the cases of
	412	sympatry from western Madagascar, where locally restricted species co-occur
	413	with the habitat generalist M. murinus (Kamilar, Blanco & Muldoon, 2016;
	414

409 lehilahytsara and M. mittermeieri are more widely distributed. Consequently, the 410 cases of sympatry between M.

. This moderate sensitivity to altitude may have 440 limited the colonization potential of M. sp. #3 southwards. These complex 441 patterns demonstrate that the biogeography and phylogeography of mouse 442 lemurs in this region of Madagascar are still not completely understood and

		Schüßler et al. page 20
	450	(between Riamalandy; Figure 1; (Weisrock et al., 2010) and Tsinjoarivo (430 km
	451	further south; (Yoder et al., 2016)). Besides in the IRS of Ambavala, the other
	452	lowland regions are always inhabited by different mouse lemur species (from
	453	north to south: M. simmonsi (Louis et al., 2006), M. gerpi (Radespiel et al.,
	454	2012), M. marohita (Rasoloarison et al., 2013)). Phylogenomic analyses
	455	indicated, that M. lehilahytsara and M. mittermeieri should rather be considered
	456	a single widely distributed species with extensive population structure, which is
	457 458 459	most likely driven by isolation-by-distance between subpopulations (Figure 2; F Poelstra et al., 2020). Under these circumstances, allopatry of southern M. o r lehilahytsara populations with lowland taxa, but sympatry of more northern
	460 461 462	P populations with lowland species (i.e., M. sp. #3 (at Ambavala) and M. e macarthurii (at Anjiahely)) requires further investigation concerning the relative e r importance of competitive exclusion, different habitat preferences or
	463 464 465	R distributional barriers, all of which have remained unexplored so far. e v i e w Morphometric differences among mouse lemurs
	466
	443	should be re-evaluated.
	444	Biogeographic patterns in the region are further complicated by our unexpected
	445	finding that the previously regarded "highland specialist" M. lehilahytsara
	446	(Radespiel et al., 2012) also occurs in the lowland rainforests around the village
	447	of Ambavala (233-462 m a.s.l). Prior to our study, this species had never been
	448	observed at altitudes below 800 m a.s.l., although it is known to occur in an
	449	extensive stretch of highland forests in central and northeastern Madagascar

Table 3

 3

	). Even sister taxa could be

Table 2 :

 2 Occurrence locations, altitudinal range (m a.s.l.) and estimated Extent 1091 of Occurrence (EOO in km² as defined by the IUCN, 2012; see Figure 1) of 1092 Microcebus spp. in northeastern Madagascar. Combined EOO for M.

	Schüßler et al. page 32

743

relative, M. macarthurii, M. jonahi is longer, has a shorter tail, wider ears, a 744 larger head width and a shorter head length. In addition, M. jonahi can be 745 differentiated from M. macarthurii by its ventral coloration which is rather whitish 746 (Figure

6

), but distinctly yellowish orange in M. macarthurii

(Radespiel et al., 747 2008; Radespiel & Raveloson, unpubl. data)

.

748

Moreover, it can be easily distinguished from the sympatric, small-bodied M.

749 lehilahytsara (at Ambavala) by its higher body mass, larger body size and 750 longer tail length. Finally, M. jonahi can be differentiated from its southern 751 geographical neighbor, M. simmonsi, by its shorter ear length and its larger 752 inter-and intra-orbital distances. M. jonahi could be unambiguously 770 English name: Jonah's mouse lemur, French name: Microcèbe de Jonah, 771 German name: Jonah's Mausmaki. 774 This study was conducted under the research permit No. 775 197/17/MEEF/SG/DGF/DSAP/SCB.Re (DS), 780 (Direction Régional de l'Environnement, de l'Ecologie et de Forêts). Our biggest 781 thanks go to the local authorities who allowed us to conduct our study in their 789 Misandeau) for their valuable help during sample collection. We warmly thank 790 the many local guides and cooks for sharing their incomparable expertise and 791 help in the field, misaotra anareo jiaby. We finally would like to thank the editor 792 and the two referees for their positive and constructive comments that helped to 793 clarify the manuscript and improve its quality.

794 Funding was granted by the Bauer Foundation and the Zempelin Foundation of 795 the "Deutsches Stiftungszentrum" under grant no. T237/22985/2012/kg and 796 T0214/32083/2018/sm to DS, Duke Tropical Conservation Initiative Grant to 797 ADY, and Duke Lemur Center/SAVA Conservation research funds to MBB, the 798 School of Animal Biology at The University of Western Australia to AM, the 799 Fundação para a Ciência e a Tecnologia, Portugal (PTDC/BIA-804 Ciência, Portugal to LC and JS, the ERA-NET BiodivERsA project: 805 INFRAGECO (Inference, Fragmentation, Genomics, and Conservation, ANR-904 monomorphic primate. Proceedings of the Royal Society B: Biological 905 Sciences, 275(1643), 1635-1644. https://doi.org/10.1098/rspb.2008.0200 906 Lei, R., Frasier, C.L., Hawkins, M.T., Engberg, S.E., Bailey, C.A., Johnson, 907 S.E., … Louis, E.E. (2016). Phylogenomic reconstruction of sportive lemurs 908 (genus Lepilemur) recovered from mitogenomes with inferences for 909 Madagascar biogeography. Journal of Heredity, 108(2), 107-119.

910 https://doi.org/10.1093/jhered/esw072 911 Louis, E.E., Coles, M.S., Andriantompohavana, R., Sommer, J.A., Engberg, 912 S.E., Zaonarivelo, J.R., … Brenneman, R.A. (2006). Revision of the mouse 949 in cryptic primates: microendemic mouse lemurs and the multispecies 950 coalescent. BioRxiv. https://doi.org/10.1101/742361 951 R Core Team (2019). R: A Language and Environment for Statistical 952 Computing. R Foundation for Statistical Computing, Vienna, Austria. URL 953 https://www.R-project.org/ 954 RStudio Team (2016). RStudio: Integrated Development Environment for R. 955 RStudio Inc., Boston MA, USA. URL: http://www.rstudio.com/ 956 Radespiel, U. (2016). Can behavioral ecology help to understand the divergent 957 geographic range size of mouse lemurs? In S.M. Lehman, U. Radespiel & 958 E. Zimmermann (Editors), The Dwarf and Mouse Lemurs of Madagascar: 959 Biology, Behavior and Conservation Biogeography of the Cheirogaleidae. 960 (pp. 498-519). Cambridge, UK: Cambridge University Press 963 Exceptional diversity of mouse lemurs (Microcebus spp.) in the Makira 972 Rakotondravony, R. & Radespiel, U. (2009). Varying patterns of coexistence of 973 two mouse lemur species (Microcebus ravelobensis and M. murinus) in a 974 heterogeneous landscape. American Journal of Primatology, 71(11), 928-975 938. https://doi.org/10.1002/ajp.20732 976 Randrianambinina, B. (2001). Contribution à l'etude comparative de 977 l'ecoethologie de deux microcebes rouges de Madagascar: Microcebus 978 ravelobensis

[START_REF] Zimmermann | Sympatric mouse lemurs in north-west Madagascar: A new rufous 1057 mouse lemur species (Microcebus ravelobensis)[END_REF]) Microcebus rufus (Lesson, 1840)

. 979 Ph.D. Thesis, University of Antananarivo 980 Randrianambinina, B., Rakotondravony, D., Radespiel, U. & Zimmermann, E. 981 (2003). Seasonal changes in general activity, body mass and reproduction

Table 3 :

 3 Morphometric measurements of Microcebus spp. in northeastern Madagascar (mean ± standard deviation).

	1097	Comparisons are based on one-way ANOVA (P < 0.001 for all parameters) and grouping (letters after values) according to Tukey
	1098	post-hoc tests. Sample sizes per site for M. sp. #3 are N = 5 at Ambavala, N = 7 at Mananara-Nord NP, N = 6 at Antanambe. §:
	1099	N = 11 for M. sp. #3.
		Variables	M. sp. #3		M. macarthurii		M. simmonsi		M. mittermeieri		M. lehilahytsara		M. lehilahytsara
		(in mm)	(N = 18, at Ambavala,	(N = 14, at Anjiahely, this	(N = 8, at Ambodiriana, this	(N = 22, at Anjiahely,		(N = 29, at Mantadia		(N = 3, at Ambavala,
			Mananara-Nord NP and	study and Radespiel et al.,		study)		Radespiel et al., 2008)		NP,			this study)
			Antanambe, this study)		2008)										Randrinambinina,
																	2001)
		Ear length	17.2 ± 0.8	a	18.0	±	0.8	a,c	19.5	±	2.7	b,c	18.3	±	1.4	a,c	20.0	±	1.1	b	17.9 ± 0.2
		Ear width §	12.7 ± 1.0	a	10.6	±	1.3	b	12.6	±	0.9	a	12.1	±	0.8	a	12.3	±	0.9	a	11.8 ± 1.1
		Head length	36.2 ± 1.4	a	37.5	±	1.3	b	35.3	±	0.9	a,c	34.3	±	0.8	c	33.2	±	1.0	d	35.2 ± 1.3
		Head width	21.5 ± 0.9	a	20.0	±	1.5	b	20.4	±	0.2	a,c	19.3	±	0.7	b,c	18.6	±	1.0	c	20.0 ± 1.2
		Snout length	10.7 ± 1.2	a	9.6	±	1.4	a	10.0	±	1.8	a	7.8	±	0.9	b	4.9	±	0.6	c	9.6 ± 0.7
		Interorbital dist. §	24.7 ± 1.1	a	23.6	±	1.5	a,b	22.6	±	0.7	b	20.6	±	1.1	c	20.1	±	0.6	c	21.6 ± 2.0
		Intraorbital dist. §	7.7 ± 0.5	a	7.1	±	0.6	a,b	6.5	±	0.6	b	6.6	±	0.7	b	5.8	±	0.5	c	7.1 ± 0.6
		Lower leg length §	39.9 ± 1.9	a	39.9	±	1.3	a	38.7	±	3.0	a	35.4	±	1.5	b	35.7	±	1.3	b	37.5 ± 0.9
		Hind foot length §	22.7 ± 1.1	a	23.3	±	1.5	a	23.4	±	2.7	a	20.2	±	1.3	b	20.3	±	1.0	b	20.7 ± 0.5
		Third toe length §	9.8 ± 1.2	a	9.5	±	0.9	a	NA	±	NA	NA	7.9	±	0.5	b	7.1	±	0.6	c	8.0 ± 0.2
		Tail length	132.7 ± 9.0	a	142.6	±	5.5	b	130.6	±	9.6	a	121.5	±	8.3	c	115.2	±	5.1	d	108.7 ± 19.7
		Body length §	93.3 ± 6.3	a	89.9	±	7.0	b	89.2	±	6.1	a,c	70.9	±	4.5	c	71.3	±	4.5	d	80.6 ± 6.4
		Body mass (g)	57.4 ± 5.6	a	61.1	±	7.8	a	51.8	±	9.1	a,b	45.1	±	6.2	b	43.8	±	4.7	b	43.3 ± 5.5

Table S1 :

 S1 Morphometric and reproductive dataset provided as supplementary xlsx-file.

	Page 63 of 66	American Journal of Primatology
		A new species of mouse lemur -page 3
	Supplementary Supplementary
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Table S2 :

 S2 Coordinates and altitude of sampling locations of Microcebus spp. used in this study.

											Supplementary Table S1 for:
	Individual.ID	Location	Age	Sex	Species	weight	ear.length	ear.width	head.length	head.width	interorbital.dist	intraorbital.dist	snout.length	lower.leg.length	hind.foot.length	third.toe.length	body.length	tail.length	testes_state	female_reproduction	Date	Collector
	B-MF-BC2	Ambavala	young	F	lehilahytsara	30	15,5		10,8	32,3	19,1	19,4		4,9	8,6	33,1		20,8	8,3		79,1	111,0	NA	closed	8-Sep-2017	DS
	B-MM-B12	Ambavala	juvenile	M	lehilahytsara	19	14,4		9,3	30,4	15,5	19,8		5,9	8,1	29,6		18,4	7,9		63,2	81,0	regressed	NA	9-Sep-2017	DS
	B-MM-B23	Ambavala	adult	M	lehilahytsara	37	18,0		11,6	33,8	18,9	19,6		7,2	10,4	36,5		20,4	8,2		75,3	86,0	regressed	NA	9-Sep-2017	DS
	B-MM-B14	Ambavala	young	M	lehilahytsara	30	16,4		11,7	33,2	19,3	20,2		8,0	8,8	34,1		22,5	9,4		79,1	119,0	regressed	NA	12-Sep-2017	DS
	B-MM-C12	Ambavala	adult	M	lehilahytsara	46	18,1		10,8	36,4	21,2	23,5		6,4	9,5	38,0		20,5	7,9		78,7	122,0	regressed	NA	15-Sep-2017	DS
	B-MF-BC3	Ambavala	adult	F	lehilahytsara	47	17,7		13,0	35,4	19,9	21,7		7,6	9,0	38,0		21,3	8,0		87,7	118,0	NA	closed	19-Sep-2017	DS
	B-MM-C24	Ambavala	juvenile	M	lehilahytsara	20	13,0		10,7	29,3	17,8	17,1		6,5	5,9	25,6		18,0	8,7		57,6	76,0	regressed	NA	19-Sep-2017	DS
	B-MM-C23 F06-99 F12-99	Ambavala Mantadia Mantadia	Location juvenile M lehilahytsara adult F lehilahytsara adult F lehilahytsara	23 46 41	15,3 19,2 20,2		Longitude 10,7 30,7 17,4 12,5 33,4 19,2 13,6 34,7 18,8	18,1 20,1 20,4	Latitude 6,5 6,6 6,6 4,2 6,8 5,3	Altitude 24,4 17,8 35,1 20,7 38,2 21,9	7,5 7,6 7,6		68,3 75,0 Reference 97,0 regressed 117,0 NA 80,0 125,0 NA	NA closed closed	19-Sep-2017 5-May-1999 9-May-1999	DS BR BR
	F14-99 F32-99 F42-99 F45-99	Mantadia Mantadia Mantadia Mantadia	adult adult adult adult	F F F F	lehilahytsara lehilahytsara lehilahytsara lehilahytsara	45 45 39 46	20,4 20,0 18,1 19,8		11,2 11,9 11,4 11,6	33,8 33,8 34,9 34,1	19,2 19,2 17,5 19,1	20,0 20,1 20,4 20,3		5,5 5,0 5,6 5,8	4,1 4,3 5,1 4,3	34,8 35,1 35,8 [m a.s.l.] 19,0 21,2 21,1 37,2 22,6	7,3 7,3 7,2 7,4		70,0 75,0 70,0 70,0	110,0 115,0 120,0 115,0	NA NA NA NA	closed closed closed open	18-May-1999 4-Sep-1999 24-Oct-1999 9-Nov-1999	BR BR BR BR
	F47-99 F48-99 F49-99	Marojejy NP Mantadia adult F lehilahytsara 50 Mantadia adult F lehilahytsara 49 Mantadia adult F lehilahytsara 55	18,5 21,7 21,0	+49º 50' 21'' -14º 28' 02'' 12,0 33,2 17,5 20,0 5,8 5,1 12,7 33,8 18,9 19,9 4,7 4,8 13,0 32,3 18,2 18,7 5,5 4,9	35,9 37,2 37,5	344 20,3 20,2 18,8	7,0 6,8 Weisrock et al., 2010 70,0 115,0 NA pregnant 70,0 115,0 NA pregnant 6,4 70,0 115,0 NA pregnant	7-Dec-1999 7-Dec-1999 7-Dec-1999	BR BR BR
	M07-99 M03-99 M10-99	Anjanaharibe-Mantadia adult M lehilahytsara 43 Mantadia adult M lehilahytsara 52 Mantadia adult M lehilahytsara 37	22,1 18,5 21,2	+49º 27' 53'' -14º 47' 27'' 13,0 32,8 19,0 20,1 6,4 5,4 11,6 32,0 18,6 20,4 6,7 4,2 12,3 33,1 19,1 18,9 5,4 4,1	37,0 34,9 35,8	1056 21,7 20,1 19,9	7,8 5,9 8,0	75,0 75,0 Louis et al., 2006 125,0 regressed NA 110,0 regressed NA 67,0 120,0 regressed NA	7-May-1999 5-May-1999 9-May-1999	BR BR BR
	M02-99	Mantadia	adult	M	lehilahytsara	46	18,8		11,3	32,5	17,9	21,9		5,2	4,3	32,5		19,4	6,7		70,0	120,0	regressed	NA	11-Apr-1999	BR
	M04-99 M08-99 M16-99 M18-99 M31-99 M33-99 M34-99 M35-99 M36-99 M37-99 M38-99 M39-99 M40-99	Mantadia Mantadia Mantadia Mantadia Mantadia Mantadia Mantadia Mantadia Mantadia Mantadia Mantadia Mantadia Mantadia	Sud SR Anjiahely Ambavala adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara adult M lehilahytsara	38 37 39 37 48 42 43 47 46 42 46 39 43	18,8 20,8 19,6 18,7 20,6 20,6 19,5 20,2 20,0 20,0 20,8 19,4 18,9	+49° 29' 38'' -15° 24' 45'' 350-400 +49° 34' 42'' -16° 11' 03'' 233-462 F o r 11,0 33,3 19,6 19,6 5,0 5,4 37,3 21,1 12,5 35,0 18,5 19,6 6,0 5,9 36,0 20,0 11,4 33,4 19,8 20,2 5,7 5,3 36,3 18,4 11,9 32,4 18,8 18,9 6,2 4,8 34,5 19,2 F 14,7 33,0 19,4 21,3 5,0 5,6 36,9 20,8 12,3 31,5 19,1 20,2 6,2 4,5 35,4 20,8 12,3 33,5 19,8 20,0 5,8 4,2 35,9 19,9 12,8 34,1 18,8 20,5 6,1 4,8 35,3 19,5 12,0 32,9 18,0 20,2 6,4 4,9 34,5 19,0 o 11,3 30,1 18,6 20,0 5,7 5,2 36,4 20,5 r 13,8 33,3 18,2 20,4 5,7 4,2 36,4 20,7 12,0 33,7 18,0 20,4 5,9 5,3 34,1 21,0 12,8 33,7 18,6 21,0 4,9 6,2 35,6 19,7	6,5 6,9 7,6 6,8 7,0 7,9 7,8 7,8 7,8 7,0 6,8 6,6 6,2		Radespiel et al., 2008, this study 65,0 115,0 regressed NA 68,0 115,0 regressed NA 70,0 115,0 regressed NA 65,0 107,0 regressed NA 80,0 115,0 enlarged NA 75,0 115,0 enlarged NA 70,0 115,0 enlarged NA 70,0 110,0 enlarged NA 70,0 110,0 enlarged NA 70,0 115,0 enlarged NA 70,0 120,0 enlarged NA 75,0 120,0 enlarged NA This study 70,0 110,0 enlarged NA	5-May-1999 7-May-1999 30-May-1999 30-May-1999 7-Aug-1999 7-Sep-1999 7-Sep-1999 7-Sep-1999 7-Sep-1999 7-Sep-1999 7-Sep-1999 24-Sep-1999 24-Sep-1999	BR BR BR BR BR BR BR BR BR BR BR BR BR
	M41-99 M43-99 M44-99 M20-99 F01-19 F11-99 M17-99 M05-99 M09-99 M13-19 M15-99 M19-99 M27-99 M28-99 M29-99 M30-99 W04-06	Mananara-Nord NP Antanambe Riamalandy Ambodiriana Mantadia adult M lehilahytsara 45 Mantadia adult M lehilahytsara 35 Mantadia adult M lehilahytsara 40 Mantadia adult M lehilahytsara NA Mantadia juvenile F lehilahytsara 20 Mantadia young F lehilahytsara 33 Mantadia young M lehilahytsara 30 Mantadia young M lehilahytsara 30 Mantadia young M lehilahytsara 36 Mantadia young M lehilahytsara 30 Mantadia young M lehilahytsara 32 Mantadia young M lehilahytsara 32 Mantadia young M lehilahytsara 33 Mantadia young M lehilahytsara 34 Mantadia young M lehilahytsara 31 Mantadia young M lehilahytsara 33 Anjiahely adult F macarthurii 50	21,7 20,6 18,5 21,9 NA NA NA NA NA NA NA NA NA NA NA NA 18,7	+49° 47' 41'' -16° 18' 18'' 277-377 +49° 48' 10'' -16° 27' 32'' 42-273 +48º 48' 54'' -16º 17' 06'' 833 +49° 42' 05'' -16° 40' 27'' 42-222 P e e r 13,5 33,8 15,6 20,0 5,7 5,3 36,6 21,3 12,4 31,6 16,7 19,6 6,3 5,0 33,5 20,3 11,3 32,9 18,2 19,9 6,0 4,6 34,8 18,8 13,5 33,7 21,0 20,0 5,7 4,6 35,9 19,6 NA NA NA NA NA NA NA NA P NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA e NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA e NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA NA r 10,2 36,5 19,7 22,0 6,9 8,4 39,7 21,4	This study This study Weisrock et al., 2010 This study 6,5 75,0 116,0 enlarged NA 7,3 60,0 115,0 enlarged NA 5,7 70,0 100,0 enlarged NA 7,2 80,0 120,0 enlarged NA NA NA NA NA closed NA NA NA NA closed NA NA NA regressed NA NA NA NA regressed NA NA NA NA regressed NA NA NA NA regressed NA NA NA NA regressed NA NA NA NA regressed NA NA NA NA regressed NA NA NA NA regressed NA NA NA NA regressed NA NA NA NA regressed NA 8,1 75,0 151,0 NA swollen	26-Sep-1999 7-Nov-1999 9-Nov-1999 31-May-1999 25-Mar-1999 9-May-1999 30-May-1999 5-May-1999 7-May-1999 18-May-1999 18-May-1999 31-May-1999 9-Aug-1999 5-Aug-1999 7-Aug-1999 7-Aug-1999 9-Nov-2006	BR BR BR BR BR BR BR BR BR BR BR BR BR BR BR BR DWR
	M03-06 W01-07 M01-06 M05-08 M0608 M07-08 M08-08 F04-11 M03-11 M01-13 F02-13 F03-13 F04-13 W06-06 M01-06leng W02-06 M07-06 M05-06 M02-07 M03-07 W04-07 M05-07 M06-07 M07-07 W08-07 M09-07 W10-07	Tampolo adult M macarthurii Zahamena NP Anjiahely 68 Betampona SNR Mantadia NP Ankofabe Ambohitantely Tsinjoarivo Anjiahely adult F macarthurii 53 Anjiahely adult M macarthurii 58 Anjiahely adult M macarthurii 72 Anjiahely adult M macarthurii 68 Anjiahely adult M macarthurii 62 Anjiahely adult M macarthurii 55 Anjiahely adult F macarthurii 72 Anjiahely adult M macarthurii 61 Anjiahely adult M macarthurii 70 Anjiahely adult F macarthurii 61 Anjiahely adult F macarthurii 51 Anjiahely adult F macarthurii 55 Anjiahely adult F mittermeieri 51 Anjiahely adult M mittermeieri 50 Anjiahely adult F mittermeieri 49 Anjiahely adult M mittermeieri 40 Anjiahely adult M mittermeieri 40 Anjiahely adult M mittermeieri 49 Anjiahely adult M mittermeieri 41 Anjiahely adult F mittermeieri 49 Anjiahely adult M mittermeieri 38 Anjiahely adult M mittermeieri 43 Anjiahely adult M mittermeieri 48 Anjiahely adult F mittermeieri 39 Anjiahely adult M mittermeieri 40 Anjiahely adult F mittermeieri 38	18,0 19,5 18,8 17,5 17,2 17,6 17,2 19,4 17,9 17,1 17,9 17,7 17,5 19,1 16,5 19,7 17,7 19,2 20,0 18,7 18,9 18,2 19,9 17,8 18,7 17,5 18,6	+49º 25' 00'' -17º 16' 60'' 12,0 37,5 19,8 23,3 6,8 8,0 +48º 44' 30'' -17º 29' 13'' +49º 12' 12'' -17º 55' 52'' +48º 25' 29'' -18º 48' 29'' +47° 11' 15'' -18° 05' 27'' +47° 16' 26'' -18° 28' 33'' +47° 41' 43'' -19° 40' 22'' R e v 40,2 956 19 21,6 300 969 1459 1271 1552 i e w 11,1 35,1 19,7 21,6 6,4 8,3 38,7 23,5 10,2 36,6 18,4 21,1 6,2 8,1 38,2 21,5 8,5 39,7 22,4 24,4 8,0 11,3 40,8 25,5 9,6 37,2 20,1 24,7 6,7 10,0 41,6 23,2 R 10,3 39,0 21,3 25,6 7,8 10,8 40,6 25,4 9,6 37,1 22,3 25,0 7,6 9,3 39,9 22,5 10,3 38,7 22,0 23,8 8,1 12,0 42,7 24,5 9,1 39,2 20,2 22,7 6,7 9,8 39,9 23,8 e 11,8 37,6 17,8 26,6 7,2 NA 38,4 NA 11,3 37,4 19,6 23,7 7,5 NA 39,6 NA 11,5 35,6 18,4 23,1 6,7 NA 37,8 NA 13,3 37,4 18,9 23,2 7,1 NA 39,8 NA v 12,2 34,0 19,2 20,7 5,4 7,0 35,2 20,4 12,1 34,3 NA 22,4 NA 9,3 38,7 21,9 i 13,4 35,4 18,7 21,4 6,7 8,8 36,2 22,9 12,7 32,5 18,8 22,6 6,4 6,3 33,8 20,0 12,9 33,0 17,9 20,3 6,2 7,7 35,5 19,8 11,8 34,7 18,8 20,6 6,6 8,0 36,6 19,4 e 10,8 33,3 19,3 20,9 6,0 6,4 34,1 19,3 11,0 34,8 20,0 20,2 6,6 7,6 36,4 20,8 11,8 34,4 20,0 19,8 6,5 7,2 34,9 20,0 12,4 34,2 18,1 19,0 6,4 8,9 35,6 20,7 12,8 34,5 19,7 20,4 7,9 6,9 35,3 20,6 w 11,9 34,5 19,9 20,9 5,6 7,8 34,2 21,0 11,6 35,3 19,0 20,9 6,1 7,7 35,8 19,2 11,0 33,8 20,0 20,5 7,0 8,7 34,6 21,0	8,2 Kappeler et al., 2005 85,0 142,0 enlarged NA Louis et al., 2006 Louis et al., 2006 Louis et al., 2006 Yoder et al., 2016 Weisrock et al., 2010 8,2 85,0 NA NA closed 8,8 82,0 142,0 enlarged NA 10,0 95,0 143,0 NA NA 9,8 94,0 144,0 enlarged NA 9,9 94,0 138,0 enlarged NA 10,0 96,0 145,0 enlarged NA 10,9 99,0 146,0 NA closed 9,8 91,0 134,0 enlarged NA 10,3 96,6 141,8 enlarged NA 9,3 94,9 141,5 NA closed NA 82,8 133,1 NA closed 10,1 88,3 151,9 NA closed 6,9 75,0 125,0 NA open 8,7 68,0 116,0 enlarged NA 7,7 80,0 119,0 NA closed 7,6 72,0 107,0 enlarged NA 7,5 73,0 121,0 enlarged NA 7,5 70,0 119,0 enlarged NA 7,6 68,0 115,0 enlarged NA 8,4 70,0 135,0 NA closed 8,0 65,0 134,0 enlarged NA 7,9 72,0 115,0 enlarged NA 7,1 70,0 125,0 enlarged NA 8,0 70,0 120,0 NA closed 7,5 67,0 128,0 enlarged NA Yoder et al., 2016 8,0 65,0 127,0 NA closed	9-Nov-2006 4-Sep-2007 8-Nov-2006 12-Dec-2008 13-Dec-2008 13-Dec-2008 13-Dec-2008 22-Nov-2011 22-Nov-2011 25-Oct-2013 25-Oct-2013 25-Oct-2008 26-Oct-2008 9-Nov-2006 13-Dec-2006 8-Nov-2006 12-Nov-2006 9-Nov-2006 4-Sep-2007 5-Sep-2007 5-Sep-2007 5-Sep-2007 5-Sep-2007 5-Sep-2007 5-Sep-2007 6-Sep-2007 6-Sep-2007	DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR DWR
	M11-07	Anjiahely	adult	M	mittermeieri	52	20,0		12,0	33,4	19,0	20,8		6,2	8,0	34,4		19,6	7,8		75,0	122,0	enlarged	NA	6-Sep-2007	DWR
	M12-07	Anjiahely	adult	M	mittermeieri	48	18,8		11,8	34,6	19,8	20,4		7,0	8,3	35,2		20,0	7,9		75,0	125,0	enlarged	NA	6-Sep-2007	DWR
	M13-07	Anjiahely	adult	M	mittermeieri	58	19,8		13,8	35,3	21,0	21,6		6,5	8,3	37,9		20,1	7,7		80,0	135,0	enlarged	NA	6-Sep-2007	DWR
	M14-07	Anjiahely	adult	M	mittermeieri	46	16,5		11,0	35,7	19,0	20,2		7,1	9,1	37,1		20,0	8,0		65,0	125,0	enlarged	NA	6-Sep-2007	DWR
	M15-07	Anjiahely	adult	M	mittermeieri	42	15,0		12,1	34,7	19,0	18,7		7,1	8,2	33,7		18,1	8,6		70,0	100,0	enlarged	NA	6-Sep-2007	DWR
	W16-07	Anjiahely	adult	F	mittermeieri	34	16,0		12,1	33,4	19,2	18,6		6,8	7,0	32,7		18,0	8,2		65,0	120,0	NA	closed	6-Sep-2007	DWR
	W17-07	Anjiahely	adult	F	mittermeieri	42	17,1		11,3	33,7	20,0	19,6		5,8	6,6	33,4		18,2	8,6		73,0	118,0	NA	closed	6-Sep-2007	DWR
	W08-06	Anjiahely	adult	F	mittermeieri	55	18,1		13,4	34,4	19,7	22,2		8,4	7,9	36,7		22,8	8,7		NA	123,0	NA	open	17-Nov-2006	DWR
	M01-08	Anjiahely	juvenile	M	mittermeieri	20	11,3		9,8	27,3	18,0	15,4		4,0	9,0	25,9		17,2	6,3		51,0	85,0	regressed	NA	11-Dec-2008	DWR
	M02-08	Anjiahely	juvenile	M	mittermeieri	20	15,7		9,2	31,1	17,5	18,0		6,4	7,9	28,5		20,0	7,7		56,0	100,7	regressed	NA	11-Dec-2008	DWR
	W03-08	Anjiahely	juvenile	F	mittermeieri	22	14,9		8,7	32,6	18,0	18,7		7,6	7,9	31,1		20,8	8,2		74,8	93,5	NA	closed	11-Dec-2008	DWR
	W04-08	Anjiahely	juvenile	F	mittermeieri	24	15,3		7,2	31,8	18,5	18,7		6,4	8,0	31,3		20,8	8,3		58,8	99,7	NA	closed	11-Dec-2008	DWR
	A1	Ambodiriana	adult	F	simmonsi	50	NA		12,9	35,5	20,4	22,2		7,5	11,9	NA		23,3	NA		NA	148,0	NA	closed	18-Jun-2014	AM
	A2	Ambodiriana	adult	F	simmonsi	46	NA		12,4	33,7	20,7	22,1		6,0	10,5	NA		29,8	NA		NA	121,0	NA	closed	18-Jun-2014	AM
	A3	Ambodiriana	adult	F	simmonsi	43	15,5		11,5	35,1	20,2	22,2		6,8	11,0	37,8		22,7	NA		80,0	130,0	NA	closed	21-Jun-2014	AM
	A4	Ambodiriana	adult	M	simmonsi	70	20,7		12,8	36,8	20,5	23,0		6,4	11,5	41,3		22,3	NA		95,0	138,0	enlarged	NA	2-Jul-2014	AM
	A5	Ambodiriana	adult	M	simmonsi	43	16,7		11,5	34,6	20,1	21,9		5,7	7,3	33,2		20,6	NA		85,0	120,0	enlarged	NA	2-Jul-2014	AM
	A6	Ambodiriana	adult	M	simmonsi	54	21,0		12,7	35,1	20,7	22,4		6,2	7,4	39,0		22,7	NA		93,0	123,0	enlarged	NA	17-Jul-2014	AM
	A7	Ambodiriana	adult	M	simmonsi	58	22,0		14,4	36,0	20,2	23,7		7,1	10,2	40,1		23,1	NA		95,0	130,0	enlarged	NA	17-Jul-2014	AM
	A8	Ambodiriana	adult	M	simmonsi	50	20,9		12,4	35,8	20,2	23,4		6,4	10,5	40,8		22,3	NA		87,0	135,0	enlarged	NA	17-Jul-2014	AM
	B-MF-BD1	Ambavala	adult	F	sp3	64	17,0		12,4	39,7	20,0	26,7		7,5	10,4	41,1		24,0	10,4		103,0	151,0	NA	closed	4-Sep-2017	DS
	B-MM-B34	Ambavala	adult	M	sp3	66	17,6		13,7	37,7	23,0	26,0		8,2	10,0	41,7		24,5	10,6		95,6	130,0	enlarged	NA	6-Sep-2017	DS
	B-MM-B13	Ambavala	adult	M	sp3	63	18,0		11,8	36,8	20,0	24,9		7,0	10,3	39,3		21,5	8,4		94,0	138,0	enlarged	NA	11-Sep-2017	DS
	B-MM-B24	Ambavala	adult	M	sp3	62	15,3		11,5	35,8	21,3	23,7		7,6	9,0	37,4		20,8	9,9		92,6	135,0	enlarged	NA	13-Sep-2017	DS
	B-MF-BC1	Ambavala	adult	F	sp3	64	18,0		12,5	36,3	23,1	23,8		8,2	10,6	39,2		23,0	8,9		96,3	132,0	NA	closed	17-Sep-2017	DS
	A-MM-A12	Antanambe	adult	M	sp3	50	18,7		13,5	36,7	21,4	24,9		7,3	10,1	38,1		22,5	11,6		95,9	128,0	enlarged	NA	13-Aug-2017	DS
	A-MM-A23	Antanambe	adult	M	sp3	54	18,0		13,5	35,9	21,6	23,2		7,0	11,4	38,4		22,9	10,4		94,0	137,0	enlarged	NA	15-Aug-2017	DS
	A-MM-A34	Antanambe	adult	M	sp3	63	17,6		14,7	34,5	22,0	24,7		8,1	11,0	44,0		23,0	8,7		100,1	133,0	enlarged	NA	17-Aug-2017	DS
	A-MM-A24	Antanambe	adult	M	sp3	53	17,0		12,1	32,8	21,3	25,4		8,1	9,1	40,1		22,6	10,9		82,8	146,0	enlarged	NA	22-Aug-2017	DS
	A-MM-A13	Antanambe	adult	M	sp3	52	18,5		12,1	37,4	20,7	24,0		8,1	9,9	39,8		23,0	7,9		83,3	146,0	enlarged	NA	24-Aug-2017	DS
	A-MF-AB1	Antanambe	adult	F	sp3	56	17,0		12,4	36,3	20,1	24,2		7,5	9,1	39,4		21,5	10,4		88,7	135,0	NA	closed	26-Aug-2017	DS
	MBB 019	Mananara_Nord adult	M	sp3	55	17,4		NA	36,7	21,6	NA		NA	13,8	NA		NA	NA		NA	131,9	enlarged	NA	13-Aug-2015	MBB
	MBB 020	Mananara_Nord adult	M	sp3	54	16,4		NA	35,7	21,4	NA		NA	11,1	NA		NA	NA		NA	131,0	enlarged	NA	15-Aug-2015	MBB
	MBB 021	Mananara_Nord adult	M	sp3	54	16,5		NA	35,4	21,4	NA		NA	10,0	NA		NA	NA		NA	131,0	enlarged	NA	15-Aug-2015	MBB
	MBB 022	Mananara_Nord young	F	sp3	46	16,3		NA	36,5	22,1	NA		NA	11,7	NA		NA	NA		NA	122,0	NA	closed	16-Aug-2015	MBB
	MBB 023	Mananara_Nord young	M	sp3	46	17,6		NA	35,2	21,6	NA		NA	11,2	NA		NA	NA		NA	119,0	enlarged	NA	18-Aug-2015	MBB
	MBB 024	Mananara_Nord young	F	sp3	47	15,4		NA	36,4	21,8	NA		NA	11,6	NA		NA	NA		NA	134,0	NA	closed	18-Aug-2015	MBB
	MBB 025	Mananara_Nord adult	M	sp3	53	16,9		NA	35,5	22,6	NA		NA	11,7	NA		NA	NA		NA	121,0	enlarged	NA	21-Aug-2015	MBB
	MBB 026	Mananara_Nord adult	M	sp3	57	17,0		NA	36,7	21,3	NA		NA	12,4	NA		NA	NA		NA	115,6	enlarged	NA	22-Aug-2015	MBB
	MBB 027	Mananara_Nord young	F	sp3	46	15,4		NA	35,6	22,0	NA		NA	9,4	NA		NA	NA		NA	123,3	NA	closed	23-Aug-2015	MBB
	MBB 028	Mananara_Nord adult	M	sp3	64	17,1		NA	36,5	22,8	NA		NA	11,2	NA		NA	NA		NA	121,2	enlarged	NA	23-Aug-2015	MBB
	MBB 029	Mananara_Nord young	M	sp3	44	16,3		NA	35,9	20,9	NA		NA	11,7	NA		NA	NA		NA	127,0	enlarged	NA	23-Aug-2015	MBB
	MBB 030	Mananara_Nord young	F	sp3	42	14,2		NA	35,7	20,4	NA		NA	10,5	NA		NA	NA		NA	131,0	NA	closed	24-Aug-2015	MBB
	MBB 031	Mananara_Nord young	M	sp3	50	17,2		NA	36,4	20,5	NA		NA	11,9	NA		NA	NA		NA	128,7	enlarged	NA	24-Aug-2015	MBB
	MBB 032	Mananara_Nord adult	M	sp3	49	16,4		NA	35,5	21,6	NA		NA	11,0	NA		NA	NA		NA	126,0	enlarged	NA	24-Aug-2015	MBB

Ecology and morphology of mouse lemurs (M icrocebus spp.) in a hotspot of microendemism in northeastern Madagascar, with the description of a new species

  Dominik Schüßler, Marina B. Blanco, Jordi Salmona, Jelmer Poelstra, Jean B. Andriambeloson, Alex Miller, Blanchard Randrianambinina, David W. Rasolofoson, Jasmin Mantilla-Contreras, Lounès Chikhi, Edward E. Louis Jr., Anne D. Yoder & Ute Radespiel Corresponding author: Ute Radespiel, Institute of Zoology, University of Veterinary Medicine Hannover, Buenteweg 17, 30559 Hannover, Germany email: ute.radespiel@tiho-hannover.de Remarks: all values in mm except weight, this parameter is in g, NA: not available all measurements are according to Hafen et al. 1998 (Folia Primatologica 69:342-356) and Zimmermann et al. 1998 (Folia Primatologia 69:106-114) Reproductive state is provided for first-time capture

	Page 67 of 66	American Journal of Primatology
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		John Wiley & Sons

16-EBI3-0014 & FCT-Biodiversa/0003/2015) the LIA BEEG-B (Laboratoire

biodiversity conservation in northeastern Madagascar using remote sensing 1010 and comparative predictive modeling. Biodiversity and Conservation. 1011 https://doi.org/10.1007/s10531-020-01965-z 1012 Schüßler, D., Radespiel, U., Ratsimbazafy, J.H. & Mantilla-Contreras, J. (2018). 1013 Lemurs in a dying forest: Factors influencing lemur diversity and distribution 1014 in forest remnants of north-eastern Madagascar. Biological Conservation, 1015 228, 17-26. https://doi.org/10.1016/j.biocon.2018.10.008 1016 Sgarlata, G. M., Salmona, J., Le Pors, B., Rasolondraibe, E., Jan, F., 1017 Ralantoharijaona, T., ... & Chikhi, L. (2019). Genetic and morphological 1018 diversity of mouse lemurs (Microcebus spp.) in northern Madagascar: The 1019 discovery of a putative new species? American Journal of Primatology, 1020 e23070. https://doi.org/10.1002/ajp.23070 1021 Thorén, S., Linnenbrink, M. & Radespiel, U. (2011). Different competitive 1022 potential in two coexisting mouse lemur species in northwestern 1023 Madagascar. American Journal of Physical Anthropology, 145(1), 156-162. 1024 https://doi.org/10.1002/ajpa.21516 1025 Venables, W.N. & Ripley, B.D. (2002). Modern Applied Statistics with S. New 1026 York: Springer, 4th Edition. URL http://www.stats.ox.ac.uk/pub/MASS4, 1027 iSBN 0-387-95457-0 1028 Vieilledent, G., Grinand, C., Rakotomalala, F.A., Ranaivosoa, R., 1029 Rakotoarijaona, J.R., Allnutt, T.F. & Achard, F. (2018). Combining global 1030 tree cover loss data with historical national forest-cover maps to look at six 1031 decades of deforestation and forest fragmentation in Madagascar. Page 45 of 66 John Wiley & Sons American Journal of Primatology Biological Conservation, 222, 189-197. 1033 https://doi.org/10.1016/j.biocon.2018.04.008 1034 Viguier, B. (2004). Functional adaptations in the craniofacial morphology of 1035 Malagasy primates: shape variations associated with gummivory in the 1036 family Cheirogaleidae. Annals of Anatomy -Anatomischer Anzeiger, 186(5-1037 6), 495-501. https://doi.org/10.1016/S0940-9602(04)80093-1 1038 Weisrock, D.W., Rasoloarison, R.M., Fiorentino, I., Ralison, J.M., Goodman, 1039 S.M., Kappeler, P.M. & Yoder, A.D. (2010). Delimiting species without 1040 nuclear monophyly in Madagascar's mouse lemurs. PLoS One, 5(3), e9883. 1041 https://doi.org/10.1371/journal.pone.0009883 1042 Wrogemann, D. & Zimmermann, E. (2001). Aspects of reproduction in the 1043 eastern rufous mouse lemur (Microcebus rufus) and their implications for 1044 captive management. Zoo Biology, 20(3), 157-167. 1045 https://doi.org/10.1002/zoo.1017 1046 Yoder, A.D., Campbell, C.R., Blanco, M.B., dos Reis, M., Ganzhorn, J.U., 1047 Goodman, S.M., … Weisrock, D.W. (2016). Geogenetic patterns in mouse 1048 lemurs (genus Microcebus) reveal the ghosts of Madagascar's forests past. 1049 Proceedings of the National Academy of Sciences, 113(29), 8049-8056.

1050 https://doi.org/10.1073/pnas.1601081113 1051 Yoder, A.D., Rasoloarison, R.M., Goodman, S.M., Irwin, J.A., Atsalis, S., 1052 Ravosa, M.J. & Ganzhorn, J.U. (2000). Remarkable species diversity in 1053 Malagasy mouse lemurs (Primates, Microcebus). Proceedings of the 1054 National Academy of Sciences, 97(21), 11325-11330. Page 46 of 66 John Wiley & Sons American Journal of Primatology

M. lehilahytsara


Francis Hindle

Lotta Kuuliala

Meriem Mouelhi

Arnaud Cuisset

Cédric Bray

Mathias Vanwolleghem

Frank Devlieghere

Gaël Mouret

Robin Bocquet

The minimization of food wastage is an important challenge for today's society. Microbial activity during the storage of foodstuffs packed under a protective atmosphere, will lead to the generation of a complex mixture of trace gases that may be used as food spoilage indicators. THz waves have been used to monitor the generation of hydrogen sulfide in the headspace of a sealed plastic tray containing a salmon fillet portion.

I. INTRODUCTION

UR capacity to determine degree of spoilage of a given product is critical to assure its quality and economical viability [START_REF] Gustavson | Global Food Losses and Food Waste[END_REF]. Spoilage is typically due to microbial activity, and leads to the production of numerous volatile compounds. Trace quantities of these molecules accumulate in the package headspace resulting in unpleasant off-odors that eventually cause consumer rejection. THz spectroscopy has great potential for the analysis of complex gas mixtures [START_REF] Bigourd | Detection and quantification of multiple molecular species in mainstream cigarette smoke by continuous-wave terahertz spectroscopy[END_REF]. We have demonstrated THz spectroscopy for the monitoring of food spoilage by measuring H2S levels in the headspace of Atlantic salmon (Salmo salar) packaged under 100 % N2. The approach was validated by comparison with selective-ion flow-tube mass spectrometry (SIFT-MS).

II. RESULTS

Thirty samples of salmon fillets were prepared and packed in plastic trays under a pure nitrogen environment before being sealed by a top film. The samples were stored at 4.0°C for a duration of 1, 3, 5, 7, 9, 11 or 13 days before being analyzed simultaneously by SIFT-MS and THz spectroscopy. A highresolution THz spectrometer constructed using such an amplified frequency multiplication chain [START_REF] Drouin | Application of cascaded frequency multiplication to molecular spectroscopy[END_REF] was developed to probe the H2S line at 611.441 GHz with a line strength of 4.18 x 10 -21 (cm -1 /(molecule.cm -2 )). The use of a calibrated gas sample and a model for the Voigt regime measured in FM mode allowed quantitative data to be extracted. Figure 1 shows the good agreement between the THz and SIFT-MS measurements. The concentration as a function of time for both techniques indicates for a given time the sample-to-sample variation is larger than the measurement uncertainties. The uncertainty of the THz measurement was established at 100 ppb [START_REF] Hindle | Monitoring of food spoilage by high resolution THz analysis[END_REF].

Fig. 1 .

 1 Fig. 1. Left pane: concentration of H2S measured by THz spectroscopy as a function of concentration measured by SIFT-MS for each sample Right pane: measured concentration values as a function of storage time for each sample. Red bars THz, black bars SIFT-MS. Length of bars (red THz, black SIFT-MS) indicate the measurement uncertainty.