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Shrews (Eulipotyphla, Soricidae) of Guatemala Musarañas (Eulipotyphla, Soricidae) De Guatemala

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Shrews (Eulipotyphla, Soricidae) of Guatemala Musarañas (Eulipotyphla, Soricidae) De Guatemala 19 Shrews (Eulipotyphla, Soricidae) of Guatemala Musarañas (Eulipotyphla, Soricidae) de Guatemala he family Soricidae or soricids (shrews) is a diverse group of small mammals found T throughout much of the world, occurring on every continent except Antarctica and Australia. They can be active at any time of the day, possi- bly because of a relatively high metabolism that re- quires an almost constant intake of food. Their diet consists mostly of invertebrates, but may include 1,* 2 occasionally small vertebrates, carrion, and vegeta- John O. Matson and Neal Woodman tion (Churchfield, 1990). 1Department of Biological Sciences, San Jose State In Guatemala, shrews occur primarily in cool University, San Jose, California, USA. moist montane forests; however, a few are found in lowland forests. In general, shrews of the genus Sorex 2United States Geological Survey, Patuxent Wildlife are restricted to montane habitats. Members of the Research Center, National Museum of Natural History, Smithsonian Institution, Washington D.C., genus Cryptotis are more generalist in their distri- USA. bution, being found in lowland as well as montane forests (Woodman et al., 2012). While they are not *Correspondence author: [email protected] arboreal, they can be found on fallen tree trunks, especially if the trees are covered with mosses. Abstract. Shrews (Soricidae) are the only members Some members of the genus Cryptotis are adapted to of the mammalian order Eulipotyphla that occur in fossorial life styles (Woodman and Stephens, 2010). Central and South America. In Guatemala, 15 species The first shrews recorded from Guatemala were have been recorded belonging to the genera Cryptotis and 9 specimens, representing 2 species, reported by Sorex, 3 of which are new and undescribed. Additionally, Gray (1843). Of these, 7 represent the genus Cryptotis 2 species are expected to be discovered in the country and the other 2 represent Sorex veraepacis Alston, based on their known distributions. Most species appear 1877. All of these early specimens were recorded as to have limited reproduction throughout the year. We being from Cobán, Alta Verapaz. However, the use review the taxonomy and natural history of these spe- of Cobán as a locality by early collectors may not cies. be accurate, because many of those specimens ac- Keywords: Cryptotis; natural history; reproductive pat- tually came from the surrounding region (Matson terns; Sorex; taxonomy. and McCarthy, 2005; McCarthy and Pérez, 2006; Woodman, 2011a; Matson and Ordóñez-Garza, 2017). The number of known species increased as more data became available (fig. 1). Reproductive patterns Reproductive patterns of Neotropical soricids are based primarily on a few scattered records over the span of decades (Woodman et al., 2012). The most complete data available for a tropical small-eared Este capítulo está sujeto a la licencia Reconocimiento-NoComercial 4.0 Internacional de Creative Commons. Para ver una copia de esta licencia, visite http://creativecommons.org/licenses/by-nc/4.0/. 20 Perspectivas de investigación sobre los mamíferos silvestres de Guatemala Figure 1. Number of species of shrews recorded from Guatemala based on data from: Gray (1843), Hall and Kelson (1959), Hall (1981), McCarthy and Pérez (2006), Woodman et al. (2012), Woodman (2018, in press) and this chapter. shrew is for Cryptotis meridensis Thomas, 1898 in the thropod abundance and availability of earthworms Venezuelan Andes. This species appears to have are positively associated with rainfall (Fragoso and the potential to reproduce throughout the year, as Lavelle, 1992; Bergallo and Magnusson, 1999), pregnant females were present in all months except so peak reproduction and lactation may coincide July and December. The proportion of pregnant fe- with higher availability of food resources. Litter males peaked in April at 23 %. Lactating females sizes (based on embryo counts) in Guatemalan and were found in all months of the year, with the high- Honduran shrews range from 1 to 2 in Sorex and est proportion (36 %) in July (Woodman and Díaz from 1 to 4 in Cryptotis, which are low compared to de Pascual, 2004). temperate species of these two genera, which can In Central America, data on reproduction reach as high as 10 young (Woodman et al., 2012; remain scarce. The most extensive dataset for Woodman, 2015). Guatemala is for Sorex ibarrai Matson and McCarthy, 2005, for which there are data based on 75 females Taxonomic Accounts representing 5 months of the year (Woodman et al., Order Eulipotyphla Waddell et al., 1999 2012). These data indicate that some females are pregnant or lactating in January, April, May, and The order Eulipotyphla comprises a subset of the July, but that a minority of the female population mammalian families that formed the tradition- is reproductively active at any given time. The rel- al, polyphyletic order Insectivora (Waddell et al., ative numbers of pregnant and lactating females 1999). The Eulipotyphla families that are consid- suggest an increased level of reproduction in April ered to form a natural group include the hedge- through July compared to January and February. hogs and gymnures (Erinaceidae), solenodons Such a pattern is tentatively supported by small- (Solenodontidae), moles (Talpidae), and shrews er datasets available from northern Neotropical (Soricidae). Only 1 of these families, the Soricidae, soricids, which also show reproductively active is present in Guatemala. In Guatemala, 2 genera females present in January, April, July, October, represent the soricids, the long-tailed shrews of the and December (Woodman et al., 2012). As a whole, genus Sorex, and the small-eared shrews of the ge- peak reproductive activity in Neotropical shrews nus Cryptotis. appears to be in April through July, with a taper- ing off toward October. If this is representative, it Family Soricidae Fischer, 1814 suggests that the majority of births and lactation The Soricidae comprises at least 25 genera and periods are timed to coincide with a rainy season more than 330 species distributed throughout that lasts from mid-May through October. Both ar- Eurasia, Africa, North and Central America, and Mamíferos terrestres pequeños 21 northern South America (Hutterer, 2005). In exter- to humid montane forests and paramos in northern nal appearance, shrews are small (head and body South America. Most species in Central America, length 35 –‍ 150 mm, mass 2 –‍ 106 g), mouse-like except C. lacandonensis Guevara et al., 2014, C. may- mammals that are often mistaken for small ro- ensis (Merriam, 1901), and C. merriami Choate, dents. Shrews typically have small pinnae that are 1970, occur above about 1000 m. They typically often concealed by fur, small eyes, and an elongat- inhabit lower montane moist forest, lower montane ed, pointed snout. They have a long, narrow, flat wet forest, montane wet forest, and montane rain skull with incomplete zygomatic arches lacking ju- forest life zones of Holdridge (1947) and may occur gals and auditory bullae. The articular condyle of in disturbed cloud forest and secondary forest. the dentary has a double articulation with the cra- In Guatemala, Cryptotis are small to medi- nium. The teeth of shrews exhibit a unique combi- um-sized shrews (length of head and body 48 –‍ 96 nation of characteristics. The deciduous dentition mm, tail 18 –‍ 34 mm, mass 4 –‍ 19 g, condylobas- is shed in utero. al length of skull or CBL 16.1 –‍ 22.8 mm, fig. 2). The Soricidae is partitioned among 3 sub- These small mammals have medium-gray to near- families: the Holarctic Soricinae, the Palearctic ly black pelage and a short- to medium-length tail Crocidurinae, and the African Myosoricinae. Only that is typically < 50 % of the length of the head members of the subfamily Soricinae occur in the and body. The tips of the cusps of the teeth are New World. The tips of the teeth of most soricines red-pigmented. There are 4 upper unicuspids that (except Megasorex and Notiosorex) have a distinctive decrease in size posteriorly, the fourth unicuspid al- red pigmentation. The large, curved first upper in- ways smaller than the third. cisor and long, procumbent first lower incisor form The species of Cryptotis are often partitioned a pincer-like foraging device. Behind the first up- taxonomically among 5 informal species groups per incisor, the anterior upper dentition (incisors, based upon external, cranial, dental, and post- canine, and anterior premolars) is comparatively cranial characteristics (Choate, 1970; Woodman, simple and undifferentiated, and these teeth are of- 1996, 2002, 2015; Woodman and Timm, 1993, ten referred to as “unicuspids”. Homologies of the 1999, 2000; Woodman et al., 2003). In Guatemala, unicuspids have been difficult to determine, and only 3 of these groups are present. for this reason, dental formulae for individual spe- The Cryptotis goldmani-group occurs from cies often disagree in the relative numbers of inci- Mexico through Honduras. Species in this group sors, canines, and premolars. Among Guatemalan are notable externally by their generally larger shrews, there are typically either 4 (Cryptotis) or 5 body size, longer pelage with little difference in col- (Sorex) unicuspids. The shrew humerus bears a teres or between dorsum and ventrum, enlarged forefeet, tubercle, a bony process mostly unique to shrews and elongate and broadened fore claws. These spe- and moles. The clavicle is long and slender. The cies are adapted for greater fossoriality than other pubic symphysis is open, so that the two halves of members of the genus, and the skeleton of the fore- the pelvis are not in contact. Shrews are sometimes limb is strongly modified. The humerus is relative- considered primitive or generalized mammals, but ly short and broad with enlarged processes for the such views ignore the unique suite of cranial, den- insertion and origin of enlarged fore limb muscles; tal, and postcranial specializations possessed by the olecranon process of the ulna is enlarged; the these small animals. bones of the forefeet are shortened and broadened, except for the distal phalanges, which are elongat- Accounts of species ed and broadened to support the enlarged claws.
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