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Hadropithecus
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<h2 id="etymology">Etymology</h2> <p>The common names that <i>Hadropithecus</i> shares with <i>Archaeolemur</i>, "<a href="/facts/Monkey_lemur/TsgXwxN1">monkey lemurs</a>" and "baboon lemurs", come from their dental and locomotor adaptations, which resemble that of modern African <a href="/facts/Baboon/qz7J3AoC">baboons</a>.<a class="footnote-ref" id="fnref:1" href="#fn:1"><sup>1</sup></a><a class="footnote-ref" id="fnref:2" href="#fn:2"><sup>2</sup></a> The genus <i>Hadropithecus</i> is derived from the <a href="/facts/Greek_language/pzDd6eIz">Greek</a> words αδρος, <i>hadros</i>, meaning "stout" or "large", and πίθηκος, <i>pithekos</i>, meaning "ape". The species name derives from the Greek root στενο-, <i>steno-</i>, meaning "narrow", and γναθος, <i>gnathos</i>, meaning "jaw" or "mouth".<a class="footnote-ref" id="fnref:3" href="#fn:3"><sup>3</sup></a> </p> <h2 id="classification-and-phylogeny">Classification and phylogeny</h2> <p><i>Hadropithecus stenognathus</i> is classified as the sole member of the genus <i>Hadropithecus</i> and belongs to the <a href="/facts/Family_(biology)/qTrn6E8E">family</a> <a href="/facts/Archaeolemuridae/TsgXwxN1">Archaeolemuridae</a>. This family in turn belongs to the infraorder <a href="/facts/Lemuriformes/b63HVq58">Lemuriformes</a>, which includes all the Malagasy lemurs.<a class="footnote-ref" id="fnref:4" href="#fn:4"><sup>4</sup></a> The species was formally described in 1899 from a <a href="/facts/Mandible/1ATEfAQa">mandible</a> (lower jaw) found at <a href="/facts/Andrahomana/Ya1M1yPw">Andrahomana</a> cave in southeastern Madagascar by <a href="/facts/Paleontologist/60n5yldd">paleontologist</a> Ludwig Lorenz von Liburnau, who thought it represented an <a href="/facts/Ape/QdTk4pko">ape</a>.<a class="footnote-ref" id="fnref:5" href="#fn:5"><sup>5</sup></a> A year later, Lorenz von Liburnau also described <i>Pithecodon sikorae</i> based on photographs of a skull, which upon further review turned out to be a juvenile version of <i>Hadropithecus stenognathus</i>. In a publication from 1902, he declared that <i>Hadrophithecus stenognathus</i> was not an ape, but a lemur.<a class="footnote-ref" id="fnref:6" href="#fn:6"><sup>6</sup></a> Over 100 years later, the rarity of its skeletal remains has made this species one of the least understood species of <a href="/facts/Subfossil_lemur/zH2drU9V">subfossil lemur</a>.<a class="footnote-ref" id="fnref:7" href="#fn:7"><sup>7</sup></a> </p> <table><tbody><tr><td><i>Hadropithecus</i> placement within the lemur phylogeny<a class="footnote-ref" id="fnref:8" href="#fn:8"><sup>8</sup></a><a class="footnote-ref" id="fnref:9" href="#fn:9"><sup>9</sup></a><a class="footnote-ref" id="fnref:10" href="#fn:10"><sup>10</sup></a></td></tr><tr><td><table><tbody><tr><td><a href="/facts/Lemur/lgLEmFnr">Lemuriformes</a></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><a href="/facts/Daubentonia/vlt4pybd">Daubentoniidae</a></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>†<a href="/facts/Koala_lemur/YDV0F4nS">Megaladapidae</a></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Lemuridae/Hg9j4thL">Lemuridae</a></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p><a href="/facts/Cheirogaleidae/ysqjVpyE">Cheirogaleidae</a></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Sportive_lemur/9WT40D6E">Lepilemuridae</a></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td>†<a href="/facts/Monkey_lemur/TsgXwxN1">Archaeolemuridae</a></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><table><tbody><tr><td>†<i><a href="/facts/Archaeolemur/UkfFo8Cd">Archaeolemur</a></i></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>†<i>Archaeolemur majori</i></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p>†<i>Archaeolemur edwardsi</i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p>†<i>Hadropithecus stenognathus</i></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><table><tbody><tr><td></td><td rowspan="2"><p>†<a href="/facts/Sloth_lemur/uMMrUfmD">Palaeopropithecidae</a></p></td></tr><tr><td></td></tr><tr><td></td><td rowspan="2"><p><a href="/facts/Indriidae/yoQLtzpL">Indriidae</a></p></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr><tr><td></td></tr></tbody></table></td></tr></tbody></table> <p>Based on similarities in their skull and teeth, it was later thought that monkey lemurs (<i>Hadropithecus</i> and <i>Archaeolemur</i>) were a sister group to the living <a href="/facts/Indri/jMIAtTpS">indriids</a> and the recently extinct <a href="/facts/Sloth_lemur/uMMrUfmD">sloth lemurs</a> (family Palaeopropithecidae).<a class="footnote-ref" id="fnref:11" href="#fn:11"><sup>11</sup></a><a class="footnote-ref" id="fnref:12" href="#fn:12"><sup>12</sup></a> However, there was some debate over whether the monkey lemurs or the sloth lemurs were more closely related to today's indriids. The monkey lemurs had skulls that more closely resembled the indriids, but their teeth were very specialized and unlike those of the indriids. The sloth lemurs, on the other hand, had teeth like the indriids, but very specialized skulls. The matter was settled with the discovery of new skeletons of <i><a href="/facts/Babakotia/VYCLbp5A">Babakotia</a></i> and <i><a href="/facts/Mesopropithecus/puUN8oP2">Mesopropithecus</a></i>, two genera of sloth lemur, both of which had indriid-like skulls and teeth.<a class="footnote-ref" id="fnref:13" href="#fn:13"><sup>13</sup></a> More recently, <a href="/facts/Postcrania/44CA9mLK">postcranial</a> remains of <i>Hadropithecus</i> found in the early 2000s prompted the suggestion that the monkey lemurs were more closely related to the <a href="/facts/Lemuridae/Hg9j4thL">lemurids</a>.<a class="footnote-ref" id="fnref:14" href="#fn:14"><sup>14</sup></a> However, <a href="/facts/DNA_sequencing/hsPsNCQW">DNA sequencing</a> has reaffirmed the sister group status of the monkey lemurs to indriids and sloth lemurs.<a class="footnote-ref" id="fnref:15" href="#fn:15"><sup>15</sup></a> </p> <h2 id="anatomy-and-physiology">Anatomy and physiology</h2> <p><i>Hadropithecus stenognathus</i> has been estimated to have weighed between 27 and 35 kg (60 and 77 lb) and to have been roughly as large as <i>Archaeolemur</i>, although more gracile.<a class="footnote-ref" id="fnref:16" href="#fn:16"><sup>16</sup></a><a class="footnote-ref" id="fnref:17" href="#fn:17"><sup>17</sup></a> Newer subfossil finds, however, suggest that <i>Hadropithecus</i> may have been more robust, and more like a <a href="/facts/Gorilla/akNvDd3d">gorilla</a> than a baboon.<a class="footnote-ref" id="fnref:18" href="#fn:18"><sup>18</sup></a> It may also have been less agile than <a href="/facts/Old_World_monkey/hrhKhgt7">Old World monkeys</a>.<a class="footnote-ref" id="fnref:19" href="#fn:19"><sup>19</sup></a> Both lemurs were <a href="/facts/Quadrupedalism/KMVYdsYf">quadrupedal</a> (walked on four legs).<a class="footnote-ref" id="fnref:20" href="#fn:20"><sup>20</sup></a> There is no evidence of <a href="/facts/Cursorial/X3I6rYVc">cursoriality</a> (adaptations specifically for running) in either species,<a class="footnote-ref" id="fnref:21" href="#fn:21"><sup>21</sup></a> and although <i>Hadropithecus</i> could have climbed trees, it lacked adaptations for leaping or <a href="/facts/Suspensory_behavior/Ma9aVERL">suspension</a>.<a class="footnote-ref" id="fnref:22" href="#fn:22"><sup>22</sup></a> </p><p>Although fewer postcranial remains have been discovered for <i>Hadropithecus</i> than for <i>Archaeolemur</i>, what has been found indicates that both were adapted for a <a href="/facts/Terrestrial_animal/AhP2mont">terrestrial</a> or semi-terrestrial lifestyle,<a class="footnote-ref" id="fnref:23" href="#fn:23"><sup>23</sup></a><a class="footnote-ref" id="fnref:24" href="#fn:24"><sup>24</sup></a><a class="footnote-ref" id="fnref:25" href="#fn:25"><sup>25</sup></a><a class="footnote-ref" id="fnref:26" href="#fn:26"><sup>26</sup></a> an unusual trait for lemurs. Both genera had short limbs and a powerful build.<a class="footnote-ref" id="fnref:27" href="#fn:27"><sup>27</sup></a> Due to its specialized dentition and likely diet, <i>Hadropithecus</i> is thought to have been the more terrestrial of the two,<a class="footnote-ref" id="fnref:28" href="#fn:28"><sup>28</sup></a> since <i>Archaeolemur</i> may have sent more time foraging and sleeping in the trees.<a class="footnote-ref" id="fnref:29" href="#fn:29"><sup>29</sup></a> Both genera also have shortened hands and feet, an adaptation for walking on the ground.<a class="footnote-ref" id="fnref:30" href="#fn:30"><sup>30</sup></a> </p><p>The face of <i>Hadropithecus</i> was shortened and adapted to heavy stress from chewing. The monkey lemurs had highly specialized teeth, but <i>Hadropithecus</i> went further by specializing in strong grinding.<a class="footnote-ref" id="fnref:31" href="#fn:31"><sup>31</sup></a> It had expanded <a href="/facts/Molar_(tooth)/gQqDtg4z">molars</a> that wore down quickly,<a class="footnote-ref" id="fnref:32" href="#fn:32"><sup>32</sup></a> much like those of <a href="/facts/Ungulate/NPtsoQmI">ungulates</a>,<a class="footnote-ref" id="fnref:33" href="#fn:33"><sup>33</sup></a> and its posterior <a href="/facts/Premolars/hSpMTu7a">premolars</a> acted like molars to extend the grinding surface.<a class="footnote-ref" id="fnref:34" href="#fn:34"><sup>34</sup></a> It also had a robust <a href="/facts/Mandible/1ATEfAQa">mandible</a> to facilitate crushing hard objects.<a class="footnote-ref" id="fnref:35" href="#fn:35"><sup>35</sup></a> Even the strepsirrhine <a href="/facts/Toothcomb/qCjKwBT1">toothcomb</a> was reduced in this species.<a class="footnote-ref" id="fnref:36" href="#fn:36"><sup>36</sup></a><a class="footnote-ref" id="fnref:37" href="#fn:37"><sup>37</sup></a><a class="footnote-ref" id="fnref:38" href="#fn:38"><sup>38</sup></a> Its <a href="/facts/Dental_formula/yMEdzfHh">dental formula</a> was 2.1.3.31.1.3.3 × 2 = 34<a class="footnote-ref" id="fnref:39" href="#fn:39"><sup>39</sup></a> </p><p>The skulls of both <i>Hadropithecus</i> and <i>Archaeolemur</i> indicate that monkey lemurs had relatively large brains compared to the other subfossil lemurs, with <i>Hadropithecus</i> having an estimated <a href="/facts/Endocranium/RJxN9qtm">endocranial</a> volume of 115 ml.<a class="footnote-ref" id="fnref:40" href="#fn:40"><sup>40</sup></a> </p> <h2 id="ecology">Ecology</h2> <p>Like all other lemurs, <i>Hadropithecus</i> was <a href="/facts/Endemic/xt4IpIsl">endemic</a> to Madagascar. Because it died out only recently and is only known from subfossil remains, it is considered to be a modern form of Malagasy lemur.<a class="footnote-ref" id="fnref:41" href="#fn:41"><sup>41</sup></a> It once ranged across the Central Plateau, South, and Southwest regions of Madagascar.<a class="footnote-ref" id="fnref:42" href="#fn:42"><sup>42</sup></a><a class="footnote-ref" id="fnref:43" href="#fn:43"><sup>43</sup></a> Within its original range, there were few other lemurs to overlap its <a href="/facts/Ecological_niche/YnhNkL2e">ecological niche</a>, and it has been shown to be the only subfossil lemur to consume both <a href="/facts/C3_carbon_fixation/HdWBsswk">C3</a> and <a href="/facts/C4_carbon_fixation/2dLTxsae">C4</a> (or <a href="/facts/CAM_photosynthesis/hnwHeIJd">CAM</a>) plants, an indication that it lived in more open habitats and had a varied diet.<a class="footnote-ref" id="fnref:44" href="#fn:44"><sup>44</sup></a><a class="footnote-ref" id="fnref:45" href="#fn:45"><sup>45</sup></a> Its physiology and <a href="/facts/Dentition/yMEdzfHh">dentition</a> suggest that it may have been much like the <a href="/facts/Gelada/GrVUDGmM">Gelada Baboon</a> in locomotion and diet,<a class="footnote-ref" id="fnref:46" href="#fn:46"><sup>46</sup></a><a class="footnote-ref" id="fnref:47" href="#fn:47"><sup>47</sup></a><a class="footnote-ref" id="fnref:48" href="#fn:48"><sup>48</sup></a> acting as a <a href="/facts/Grazing/nxF45P2c">manual grazer</a> (picking grass with the hands) since its teeth were well-adapted for grinding either grass or seeds.<a class="footnote-ref" id="fnref:49" href="#fn:49"><sup>49</sup></a><a class="footnote-ref" id="fnref:50" href="#fn:50"><sup>50</sup></a> Microwear patterns on its teeth, as well as its overly large molars, indicate it processed hard objects like nuts or seeds, making it a <a href="/facts/Seed_predation/4Cbuef4k">seed predator</a>.<a class="footnote-ref" id="fnref:51" href="#fn:51"><sup>51</sup></a><a class="footnote-ref" id="fnref:52" href="#fn:52"><sup>52</sup></a> More recent microwear analysis suggests differences between Gelada Baboons and <i>Hadropithecus</i>, indicating that this extinct lemur may not have been a grazer, but strictly a hard object processor.<a class="footnote-ref" id="fnref:53" href="#fn:53"><sup>53</sup></a> </p> <h2 id="extinction">Extinction</h2> <p>Because of the low number of subfossil finds, <i>Hadropithecus</i> is thought to have been rare,<a class="footnote-ref" id="fnref:54" href="#fn:54"><sup>54</sup></a> and it died out sooner than its sister taxon, <i>Archaeolemur</i>.<a class="footnote-ref" id="fnref:55" href="#fn:55"><sup>55</sup></a> Both disappeared shortly after the arrival of humans to the island, but being a large, specialized, terrestrial grazer, <i>Hadropithecus</i> would have faced more pressure from domestic <a href="/facts/Livestock/Quhv6GvP">livestock</a>, introduced <a href="/facts/Pigs/b5wxwdDY">pigs</a>, and spreading human populations than its more generalized cousin.<a class="footnote-ref" id="fnref:56" href="#fn:56"><sup>56</sup></a> The last known record was <a href="/facts/Radiocarbon_dating/qjNglhKP">radiocarbon dated</a> to around 444–772 <a href="/facts/Common_Era/e5DpqUoe">CE</a>.<a class="footnote-ref" id="fnref:57" href="#fn:57"><sup>57</sup></a> </p> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. pp. 37–51. ISBN 1-881173-88-7. OCLC 883321520. <a href="1-881173-88-7" target="_blank">1-881173-88-7</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Sussman, R.W. (2003). "Chapter 4: The Nocturnal Lemuriformes". Primate Ecology and Social Structure. Pearson Custom Publishing. pp. 107–148. ISBN 978-0-536-74363-3. <a href="978-0-536-74363-3" target="_blank">978-0-536-74363-3</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>Palmer, T. (1904). "Index generum mammalium: a list of the genera and families of mammals". North American Fauna. 23: 1–984. doi:10.3996/nafa.23.0001. pp. 80–81, 539, 645, 648. <a href="https://books.google.com/books?id=WdnPAAAAMAAJ" target="_blank">https://books.google.com/books?id=WdnPAAAAMAAJ</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>Nowak, R.M. (1999). "Family Archaeolemuridae: Baboon Lemurs". Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 91–92. ISBN 978-0-8018-5789-8. <a href="978-0-8018-5789-8" target="_blank">978-0-8018-5789-8</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>Lorenz von Liburnau, L. (1899). "Herr Custor Dr. Ludwig v. Lorenz berichtet über einen fossilen Anthropoiden von Madagascar". Anzeigen der Kaiserlichen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Classe, Wien. 27: 255–257. <a href="https://books.google.com/books?id=xkQUAAAAYAAJ&q=Hadropithecus&pg=PA255" target="_blank">https://books.google.com/books?id=xkQUAAAAYAAJ&q=Hadropithecus&pg=PA255</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></p></li> <li id="fn:6"><p>Lorenz von Liburnau, L. (1902). "Über Hadropithecus stenognathus Lz., nebst Bemerkungen zu einigen anderen ausgestorbenen Primaten von Madagaskar". Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Wien. 72: 243–254. <a href="https://www.biodiversitylibrary.org/item/31611#315" target="_blank">https://www.biodiversitylibrary.org/item/31611#315</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></p></li> <li id="fn:7"><p>Ryan, T.M.; Burney, D.A.; Godfrey, L.R.; Göhlich, U.B.; Jungers, W.L.; Vasey, N.; Ramilisonina; Walker, A.; Weber, G.W. (2008). "A reconstruction of the Vienna skull of Hadropithecus stenognathus" (PDF). PNAS. 105 (31): 10699–10702. Bibcode:2008PNAS..10510699R. doi:10.1073/pnas.0805195105. PMC 2488384. PMID 18663217. Retrieved 19 February 2010. <a href="http://www.pnas.org/content/105/31/10699.full.pdf" target="_blank">http://www.pnas.org/content/105/31/10699.full.pdf</a> <a href="#fnref:7" class="footnote-back-ref">↩</a></p></li> <li id="fn:8"><p>Horvath, J.; et al. (2008). "Development and application of a phylogenomic toolkit: Resolving the evolutionary history of Madagascar's lemurs" (PDF). Genome Research. 18 (3): 489–99. doi:10.1101/gr.7265208. PMC 2259113. PMID 18245770. Retrieved 2009-09-02. <a href="http://www.biology.duke.edu/yoderlab/reprints/2008Horvath_etalGR.pdf" target="_blank">http://www.biology.duke.edu/yoderlab/reprints/2008Horvath_etalGR.pdf</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></p></li> <li id="fn:9"><p>Orlando, L.; Calvignac, S.; Schnebelen, C.; Douady, C.J.; Godfrey, L.R.; Hänni, C. (2008). "DNA from extinct giant lemurs links archaeolemurids to extant indriids". BMC Evolutionary Biology. 8 (1): 121. Bibcode:2008BMCEE...8..121O. doi:10.1186/1471-2148-8-121. PMC 2386821. PMID 18442367. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2386821" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2386821</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>Godfrey, L.R.; Jungers, W.L. (2003). "Subfossil Lemurs". In Goodman, S.M.; Benstead, J.P (eds.). The Natural History of Madagascar. University of Chicago Press. pp. 1247–1252. ISBN 978-0-226-30306-2. <a href="978-0-226-30306-2" target="_blank">978-0-226-30306-2</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:11" class="footnote-back-ref">↩</a></p></li> <li id="fn:12"><p>Simons, E.L. (1997). "Chapter 6: Lemurs: Old and New". In Goodman, S.M.; Patterson, B.D (eds.). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. pp. 142–166. ISBN 978-1-56098-682-9. <a href="978-1-56098-682-9" target="_blank">978-1-56098-682-9</a> <a href="#fnref:12" class="footnote-back-ref">↩</a></p></li> <li id="fn:13"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:13" class="footnote-back-ref">↩</a></p></li> <li id="fn:14"><p>Godfrey, L.R.; Jungers, W.L.; Burney, D.A.; Vasey, N.; Ramilisonina, S.J.; Wheeler, W.; Lemelin, P.; Shapiro, L.J.; et al. (2006). "New discoveries of skeletal elements of Hadropithecus stenognathus from Andrahomana Cave, southeastern Madagascar". Journal of Human Evolution. 51 (4): 395–410. Bibcode:2006JHumE..51..395G. doi:10.1016/j.jhevol.2006.04.012. PMID 16911817. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:14" class="footnote-back-ref">↩</a></p></li> <li id="fn:15"><p>Orlando, L.; Calvignac, S.; Schnebelen, C.; Douady, C.J.; Godfrey, L.R.; Hänni, C. (2008). "DNA from extinct giant lemurs links archaeolemurids to extant indriids". BMC Evolutionary Biology. 8 (1): 121. Bibcode:2008BMCEE...8..121O. doi:10.1186/1471-2148-8-121. PMC 2386821. PMID 18442367. <a href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2386821" target="_blank">https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2386821</a> <a href="#fnref:15" class="footnote-back-ref">↩</a></p></li> <li id="fn:16"><p>Nowak, R.M. (1999). "Family Archaeolemuridae: Baboon Lemurs". Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 91–92. ISBN 978-0-8018-5789-8. <a href="978-0-8018-5789-8" target="_blank">978-0-8018-5789-8</a> <a href="#fnref:16" class="footnote-back-ref">↩</a></p></li> <li id="fn:17"><p>Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. pp. 37–51. ISBN 1-881173-88-7. OCLC 883321520. <a href="1-881173-88-7" target="_blank">1-881173-88-7</a> <a href="#fnref:17" class="footnote-back-ref">↩</a></p></li> <li id="fn:18"><p>Godfrey, L.R.; Jungers, W.L.; Schwartz, G.T. (2006). "Chapter 3: Ecology and Extinction of Madagascar's Subfossil Lemurs". In Gould, L.; Sauther, M.L (eds.). Lemurs: Ecology and Adaptation. Springer. pp. 41–64. ISBN 978-0-387-34585-7. <a href="978-0-387-34585-7" target="_blank">978-0-387-34585-7</a> <a href="#fnref:18" class="footnote-back-ref">↩</a></p></li> <li id="fn:19"><p>Ryan, T.M.; Burney, D.A.; Godfrey, L.R.; Göhlich, U.B.; Jungers, W.L.; Vasey, N.; Ramilisonina; Walker, A.; Weber, G.W. (2008). "A reconstruction of the Vienna skull of Hadropithecus stenognathus" (PDF). PNAS. 105 (31): 10699–10702. Bibcode:2008PNAS..10510699R. doi:10.1073/pnas.0805195105. PMC 2488384. PMID 18663217. Retrieved 19 February 2010. <a href="http://www.pnas.org/content/105/31/10699.full.pdf" target="_blank">http://www.pnas.org/content/105/31/10699.full.pdf</a> <a href="#fnref:19" class="footnote-back-ref">↩</a></p></li> <li id="fn:20"><p>Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. pp. 37–51. ISBN 1-881173-88-7. OCLC 883321520. <a href="1-881173-88-7" target="_blank">1-881173-88-7</a> <a href="#fnref:20" class="footnote-back-ref">↩</a></p></li> <li id="fn:21"><p>Godfrey, L.R.; Jungers, W.L.; Schwartz, G.T. (2006). "Chapter 3: Ecology and Extinction of Madagascar's Subfossil Lemurs". In Gould, L.; Sauther, M.L (eds.). Lemurs: Ecology and Adaptation. Springer. pp. 41–64. ISBN 978-0-387-34585-7. <a href="978-0-387-34585-7" target="_blank">978-0-387-34585-7</a> <a href="#fnref:21" class="footnote-back-ref">↩</a></p></li> <li id="fn:22"><p>Ryan, T.M.; Burney, D.A.; Godfrey, L.R.; Göhlich, U.B.; Jungers, W.L.; Vasey, N.; Ramilisonina; Walker, A.; Weber, G.W. (2008). "A reconstruction of the Vienna skull of Hadropithecus stenognathus" (PDF). PNAS. 105 (31): 10699–10702. Bibcode:2008PNAS..10510699R. doi:10.1073/pnas.0805195105. PMC 2488384. PMID 18663217. Retrieved 19 February 2010. <a href="http://www.pnas.org/content/105/31/10699.full.pdf" target="_blank">http://www.pnas.org/content/105/31/10699.full.pdf</a> <a href="#fnref:22" class="footnote-back-ref">↩</a></p></li> <li id="fn:23"><p>Nowak, R.M. (1999). "Family Archaeolemuridae: Baboon Lemurs". Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 91–92. ISBN 978-0-8018-5789-8. <a href="978-0-8018-5789-8" target="_blank">978-0-8018-5789-8</a> <a href="#fnref:23" class="footnote-back-ref">↩</a></p></li> <li id="fn:24"><p>Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. pp. 37–51. ISBN 1-881173-88-7. OCLC 883321520. <a href="1-881173-88-7" target="_blank">1-881173-88-7</a> <a href="#fnref:24" class="footnote-back-ref">↩</a></p></li> <li id="fn:25"><p>Sussman, R.W. (2003). "Chapter 4: The Nocturnal Lemuriformes". Primate Ecology and Social Structure. Pearson Custom Publishing. pp. 107–148. ISBN 978-0-536-74363-3. <a href="978-0-536-74363-3" target="_blank">978-0-536-74363-3</a> <a href="#fnref:25" class="footnote-back-ref">↩</a></p></li> <li id="fn:26"><p>Simons, E.L. (1997). "Chapter 6: Lemurs: Old and New". In Goodman, S.M.; Patterson, B.D (eds.). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. pp. 142–166. ISBN 978-1-56098-682-9. <a href="978-1-56098-682-9" target="_blank">978-1-56098-682-9</a> <a href="#fnref:26" class="footnote-back-ref">↩</a></p></li> <li id="fn:27"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:27" class="footnote-back-ref">↩</a></p></li> <li id="fn:28"><p>Simons, E.L. (1997). "Chapter 6: Lemurs: Old and New". In Goodman, S.M.; Patterson, B.D (eds.). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. pp. 142–166. ISBN 978-1-56098-682-9. <a href="978-1-56098-682-9" target="_blank">978-1-56098-682-9</a> <a href="#fnref:28" class="footnote-back-ref">↩</a></p></li> <li id="fn:29"><p>Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. pp. 37–51. ISBN 1-881173-88-7. OCLC 883321520. <a href="1-881173-88-7" target="_blank">1-881173-88-7</a> <a href="#fnref:29" class="footnote-back-ref">↩</a></p></li> <li id="fn:30"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:30" class="footnote-back-ref">↩</a></p></li> <li id="fn:31"><p>Godfrey, L.R.; Jungers, W.L.; Reed, K.E.; Simons, E.L.; Chatrath, P.S. (1997). "Chapter 8: Subfossil Lemurs". In Goodman, S.M.; Patterson, B.D. (eds.). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. pp. 218–256. ISBN 978-1-56098-682-9. <a href="978-1-56098-682-9" target="_blank">978-1-56098-682-9</a> <a href="#fnref:31" class="footnote-back-ref">↩</a></p></li> <li id="fn:32"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:32" class="footnote-back-ref">↩</a></p></li> <li id="fn:33"><p>Nowak, R.M. (1999). "Family Archaeolemuridae: Baboon Lemurs". Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 91–92. ISBN 978-0-8018-5789-8. <a href="978-0-8018-5789-8" target="_blank">978-0-8018-5789-8</a> <a href="#fnref:33" class="footnote-back-ref">↩</a></p></li> <li id="fn:34"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:34" class="footnote-back-ref">↩</a></p></li> <li id="fn:35"><p>Godfrey, L.R.; Jungers, W.L.; Reed, K.E.; Simons, E.L.; Chatrath, P.S. (1997). "Chapter 8: Subfossil Lemurs". In Goodman, S.M.; Patterson, B.D. (eds.). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. pp. 218–256. ISBN 978-1-56098-682-9. <a href="978-1-56098-682-9" target="_blank">978-1-56098-682-9</a> <a href="#fnref:35" class="footnote-back-ref">↩</a></p></li> <li id="fn:36"><p>Nowak, R.M. (1999). "Family Archaeolemuridae: Baboon Lemurs". Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 91–92. ISBN 978-0-8018-5789-8. <a href="978-0-8018-5789-8" target="_blank">978-0-8018-5789-8</a> <a href="#fnref:36" class="footnote-back-ref">↩</a></p></li> <li id="fn:37"><p>Sussman, R.W. (2003). "Chapter 4: The Nocturnal Lemuriformes". Primate Ecology and Social Structure. Pearson Custom Publishing. pp. 107–148. ISBN 978-0-536-74363-3. <a href="978-0-536-74363-3" target="_blank">978-0-536-74363-3</a> <a href="#fnref:37" class="footnote-back-ref">↩</a></p></li> <li id="fn:38"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:38" class="footnote-back-ref">↩</a></p></li> <li id="fn:39"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:39" class="footnote-back-ref">↩</a></p></li> <li id="fn:40"><p>Ryan, T.M.; Burney, D.A.; Godfrey, L.R.; Göhlich, U.B.; Jungers, W.L.; Vasey, N.; Ramilisonina; Walker, A.; Weber, G.W. (2008). "A reconstruction of the Vienna skull of Hadropithecus stenognathus" (PDF). PNAS. 105 (31): 10699–10702. Bibcode:2008PNAS..10510699R. doi:10.1073/pnas.0805195105. PMC 2488384. PMID 18663217. Retrieved 19 February 2010. <a href="http://www.pnas.org/content/105/31/10699.full.pdf" target="_blank">http://www.pnas.org/content/105/31/10699.full.pdf</a> <a href="#fnref:40" class="footnote-back-ref">↩</a></p></li> <li id="fn:41"><p>Sussman, R.W. (2003). "Chapter 4: The Nocturnal Lemuriformes". Primate Ecology and Social Structure. Pearson Custom Publishing. pp. 107–148. ISBN 978-0-536-74363-3. <a href="978-0-536-74363-3" target="_blank">978-0-536-74363-3</a> <a href="#fnref:41" class="footnote-back-ref">↩</a></p></li> <li id="fn:42"><p>Nowak, R.M. (1999). "Family Archaeolemuridae: Baboon Lemurs". Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 91–92. ISBN 978-0-8018-5789-8. <a href="978-0-8018-5789-8" target="_blank">978-0-8018-5789-8</a> <a href="#fnref:42" class="footnote-back-ref">↩</a></p></li> <li id="fn:43"><p>Sussman, R.W. (2003). "Chapter 4: The Nocturnal Lemuriformes". Primate Ecology and Social Structure. Pearson Custom Publishing. pp. 107–148. ISBN 978-0-536-74363-3. <a href="978-0-536-74363-3" target="_blank">978-0-536-74363-3</a> <a href="#fnref:43" class="footnote-back-ref">↩</a></p></li> <li id="fn:44"><p>Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. pp. 37–51. ISBN 1-881173-88-7. OCLC 883321520. <a href="1-881173-88-7" target="_blank">1-881173-88-7</a> <a href="#fnref:44" class="footnote-back-ref">↩</a></p></li> <li id="fn:45"><p>Godfrey, L.R.; Jungers, W.L.; Schwartz, G.T. (2006). "Chapter 3: Ecology and Extinction of Madagascar's Subfossil Lemurs". In Gould, L.; Sauther, M.L (eds.). Lemurs: Ecology and Adaptation. Springer. pp. 41–64. ISBN 978-0-387-34585-7. <a href="978-0-387-34585-7" target="_blank">978-0-387-34585-7</a> <a href="#fnref:45" class="footnote-back-ref">↩</a></p></li> <li id="fn:46"><p>Nowak, R.M. (1999). "Family Archaeolemuridae: Baboon Lemurs". Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 91–92. ISBN 978-0-8018-5789-8. <a href="978-0-8018-5789-8" target="_blank">978-0-8018-5789-8</a> <a href="#fnref:46" class="footnote-back-ref">↩</a></p></li> <li id="fn:47"><p>Sussman, R.W. (2003). "Chapter 4: The Nocturnal Lemuriformes". Primate Ecology and Social Structure. Pearson Custom Publishing. pp. 107–148. ISBN 978-0-536-74363-3. <a href="978-0-536-74363-3" target="_blank">978-0-536-74363-3</a> <a href="#fnref:47" class="footnote-back-ref">↩</a></p></li> <li id="fn:48"><p>Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 33–48. ISBN 1-881173-08-9. OCLC 32480729. <a href="1-881173-08-9" target="_blank">1-881173-08-9</a> <a href="#fnref:48" class="footnote-back-ref">↩</a></p></li> <li id="fn:49"><p>Sussman, R.W. (2003). "Chapter 4: The Nocturnal Lemuriformes". Primate Ecology and Social Structure. Pearson Custom Publishing. pp. 107–148. ISBN 978-0-536-74363-3. <a href="978-0-536-74363-3" target="_blank">978-0-536-74363-3</a> <a href="#fnref:49" class="footnote-back-ref">↩</a></p></li> <li id="fn:50"><p>Simons, E.L. (1997). "Chapter 6: Lemurs: Old and New". In Goodman, S.M.; Patterson, B.D (eds.). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. pp. 142–166. ISBN 978-1-56098-682-9. <a href="978-1-56098-682-9" target="_blank">978-1-56098-682-9</a> <a href="#fnref:50" class="footnote-back-ref">↩</a></p></li> <li id="fn:51"><p>Godfrey, L.R.; Jungers, W.L.; Schwartz, G.T. (2006). "Chapter 3: Ecology and Extinction of Madagascar's Subfossil Lemurs". In Gould, L.; Sauther, M.L (eds.). Lemurs: Ecology and Adaptation. Springer. pp. 41–64. ISBN 978-0-387-34585-7. <a href="978-0-387-34585-7" target="_blank">978-0-387-34585-7</a> <a href="#fnref:51" class="footnote-back-ref">↩</a></p></li> <li id="fn:52"><p>Godfrey, L.R.; Jungers, W.L.; Reed, K.E.; Simons, E.L.; Chatrath, P.S. (1997). "Chapter 8: Subfossil Lemurs". In Goodman, S.M.; Patterson, B.D. (eds.). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. pp. 218–256. ISBN 978-1-56098-682-9. <a href="978-1-56098-682-9" target="_blank">978-1-56098-682-9</a> <a href="#fnref:52" class="footnote-back-ref">↩</a></p></li> <li id="fn:53"><p>Ryan, T.M.; Burney, D.A.; Godfrey, L.R.; Göhlich, U.B.; Jungers, W.L.; Vasey, N.; Ramilisonina; Walker, A.; Weber, G.W. (2008). "A reconstruction of the Vienna skull of Hadropithecus stenognathus" (PDF). PNAS. 105 (31): 10699–10702. Bibcode:2008PNAS..10510699R. doi:10.1073/pnas.0805195105. PMC 2488384. PMID 18663217. Retrieved 19 February 2010. <a href="http://www.pnas.org/content/105/31/10699.full.pdf" target="_blank">http://www.pnas.org/content/105/31/10699.full.pdf</a> <a href="#fnref:53" class="footnote-back-ref">↩</a></p></li> <li id="fn:54"><p>Simons, E.L. (1997). "Chapter 6: Lemurs: Old and New". In Goodman, S.M.; Patterson, B.D (eds.). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. pp. 142–166. ISBN 978-1-56098-682-9. <a href="978-1-56098-682-9" target="_blank">978-1-56098-682-9</a> <a href="#fnref:54" class="footnote-back-ref">↩</a></p></li> <li id="fn:55"><p>Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. pp. 37–51. ISBN 1-881173-88-7. OCLC 883321520. <a href="1-881173-88-7" target="_blank">1-881173-88-7</a> <a href="#fnref:55" class="footnote-back-ref">↩</a></p></li> <li id="fn:56"><p>Nowak, R.M. (1999). "Family Archaeolemuridae: Baboon Lemurs". Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 91–92. ISBN 978-0-8018-5789-8. <a href="978-0-8018-5789-8" target="_blank">978-0-8018-5789-8</a> <a href="#fnref:56" class="footnote-back-ref">↩</a></p></li> <li id="fn:57"><p>Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. pp. 37–51. ISBN 1-881173-88-7. OCLC 883321520. <a href="1-881173-88-7" target="_blank">1-881173-88-7</a> <a href="#fnref:57" class="footnote-back-ref">↩</a></p></li> </ol>