Ants (Hymenoptera: Formicidae) and termites (Blattodea: Termitoidae) in the diet of wild white-bellied pangolin (Phataginus tricuspis) in forest-savanna habitats of Cameroon.

The white-bellied pangolin Phataginus tricuspis (Rafinesque 1821) is a semiarboreal species occurring in tropical sub-Saharan Africa. It is the world's most trafficked African pangolin species based on volumes recorded in seizures. Reintroduction of confiscated live pangolins and ex-situ rearing are being explored worldwide as a conservation action. However, the husbandry of seized animals is challenging as the diet of the white-bellied pangolin is poorly known and little studied. We analyzed the stomach contents of dead white-bellied pangolins from two forest-savanna protected areas. Stomach content samples from 13 white-bellied pangolin specimens contained ~165,000 Arthropoda, mostly Hymenoptera (60.34%) and Blattodea (39.66%). Overall, we identified 39 termite and 105 ant species consumed as prey by pangolins. Individual pangolins examined had fed on a maximum of 31 ant species and 13 termite species. The termite and ant species richness varied significantly across the pangolins' last consumed meal. We recorded 24 ant genera dominated by Crematogaster (relative importance [RI] = 17.28). Out of 18 termite genera recorded, the genus Pseudacanthotermes (RI = 17.21) was the most important prey. Ten ant species were preferentially eaten by white-bellied pangolin, with Crematogaster acis being the most common prey species. Four species of termite were most frequently eaten with Pseudacanthotermes militaris being the most abundant. The mean abundance of ants and termites varied among pangolin individuals. The season did not influence the mean abundance of termites eaten by pangolin individuals. However, ant abundance in stomach contents was significantly higher in the dry season. An improved understanding of pangolin feeding behavior and prey selection may help inform conservation husbandry efforts. For example, nutritional analysis of the food eaten by wild pangolins can guide the development of nutritional diets for captive pangolins.

Digesta retention time and recovery rates of ants and termites in Chinese pangolins (Manis pentadactyla).

Pangolins are myrmecophagous mammals whose biology and ecology remain poorly studied. Termite mandibles and ant head capsules are the two primary remains found in pangolin feces. Determining the retention time of insect cuticles is important for understanding the digestive physiology of pangolins, while determining the recovery rate of termites and ants in feces is required to estimate the number of these prey items that are consumed by pangolins. In this study, the authors conducted feeding trials with captive Chinese pangolins (Manis pentadactyla). Sixty grams of the fungus-growing termite Odontotermes formosanus (18,816 individuals) and 15-20 g of the yellow crazy ant Anoplolepis gracilipes (14,400-19,200 individuals) were fed to each pangolin. After feeding, pangolin feces were collected daily for 1 week. The authors also assessed the accuracy of using chromium (III) oxide powder (Cr2 O3 ) as a proxy for determining gut passage time, as has been done in previous studies. The results showed that remaining termite mandibles and ant head capsules in feces peaked at 66 and 90 hr after feeding and their recovery rates were 0.35 ± 0.10 and 0.65 ± 0.04, respectively. In both feeding trials, the retention time of Cr2 O3 was much shorter than that of the termite mandibles and ant head capsules, indicating that Cr2 O3 is not an appropriate indicator for estimating food retention time of myrmecophagous animals. Our results revealed that the ant head capsules were preserved better in feces compared with the termite mandibles, suggesting that termites may be considerably underestimated in the feces of wild pangolins.

COMPLICATIONS ASSOCIATED WITH PREGNANCY AND PARTURITION IN AFRICAN WHITE-BELLIED PANGOLINS (PHATAGINUS TRICUSPIS).

There are no studies to date on the normal reproductive physiology of African white-bellied pangolins (Phataginus tricuspis). As a reclusive species, little is known about normal gestation, successful parturition, and potential complications during pregnancy. Ten female P. tricuspis were diagnosed as pregnant and monitored under professional care. Five developed complications during pregnancies or during parturition and are detailed in this case series. Dystocia occurred in two dams each, with malposition of a singleton fetus. Both dams were successfully treated with surgical intervention by caesarian section. Of the two individuals, one fetus was nonviable, but the other neonate survived and was reared by the dam to weaning. A third pregnant female died during pregnancy from septicemia resulting in death of the preterm fetus. The two additional dams of the five gave birth to full-term neonates. One fetus was stillborn with evidence of fetal distress, and the other died immediately after birth (perinatal death) with undetermined etiology. Based on this case series, complications associated with pregnancy occur in P. tricuspis, indicating the need for further study and close monitoring during impending parturition.

The gross reproductive morphology of the male Temminck's pangolin Smutsia temminckii (Smuts, 1832).

The Temminck's pangolin (Smutsia temminckii) is one of eight pangolin species worldwide and the only pangolin present in southern Africa. Historically, pangolins have not been able to reproduce successfully in captivity and this may be in part due to the lack of knowledge and understanding with regards to the pangolin reproductive system (anatomy, physiology, biology) in all eight species. This original study describes the gross anatomy of the male Temminck's pangolin from three adult individuals investigated. The male Temminck's pangolin presented a short, conical penis with ascrotal (internal) testes, similar to many other myrmecophagous mammals such as the aardvark (Orycteropus sp.) and anteaters (suborder: Vermilingua). However, the orientation of the penis of the Temminck's pangolin differed in that it was oriented cranioventrally, in contrast to the caudal orientation of the giant anteater. The testes were found to be bilaterally flattened with an elongate oval shape, similar to the aardvark. The specific characteristics of the reproductive tract of the male Temminck's pangolins are thought to be adaptations to their peculiar lifestyle as the male portrays characteristics that indicate adaptation to a lower basal metabolic rate and body temperature as well as to their defensive mechanism of rolling up into a ball. Our study suggests the male Temminck's pangolin reproductive anatomy is most similar and comparable to the Xenarthrans and the aardvark that display the same fossorial activities as pangolins, and the male morphology is not comparable to the phylogenetically closely-related Carnivora.

The brain of the tree pangolin (Manis tricuspis). X. The spinal cord.

The spinal cord of the tree pangolin is known to be very short compared to the overall length of the body and tail. Here, we provide a description of the tree pangolin spinal cord to determine whether the short length contributes to specific structural, and potentially functional, differences. The short spinal cord of the adult tree pangolin, at around 13 cm, terminates at the midthoracic level. Within this shortened spinal cord, we could identify six regions, which from rostral to caudal include the prebrachial, brachial, interramal, crural, postcrural, and caudal regions, with both the brachial and crural regions showing distinct swellings. The chemoarchitecture of coronal sections through these regions confirmed regional assignation, being most readily delineated by the presence of cholinergic neurons forming the intermediolateral column in the interramal region and the sacral parasympathetic nucleus in the postcrural region. The 10 laminae of Rexed were observed throughout the spinal cord and presented with an anatomical organization similar to that observed in other mammals. Despite the shortened length of the tree pangolin spinal cord, the regional and laminar anatomical organization is very similar to that observed in other mammals. This indicates that the functional aspects of the short tree pangolin spinal cord can be inferred from what is known in other mammals.

Long term monitoring of the reproductive behavior of wild Chinese pangolin (Manis pentadactyla).

Observations of Chinese pangolin (Manis pentadactyla) in the wild are extremely rare and challenging because of their nocturnal and cryptic activity patterns and low population density. The present article reported the first field observation in eastern Taiwan, from October 4, 2012 to June 16, 2016, on the reproductive behavior of the Chinese pangolin based on the monitoring of a female (LF28) using radiotelemetry and camera traps. During this period, LF28 aged from 1-4.5 years old and gave two single-births, both took place in early December, at 3 and 4 years old, respectively. We recorded the entire 157 days of the first nursing period from parturition to maternal separation. For the second infant, the gestation period was estimated to be around 150 days based on the evidence that the pregnancy started in early Jul. 2015 and the offspring was born on Dec. 9, 2015. During the entire nursing period, LF28 frequently moved the offspring from one nursing burrow to another staying various durations ranging from 1 day to more than 35 days, and almost all (= 15/16) of these burrows were located in the core (MCP75) of LF28's home range. Started from the month of parturition and lasting throughout the whole nursing period, different adult males constantly visiting the nursing burrows were recorded. Mating behavior was recorded once outside the burrow in March, which provided evidence of the occurrence of post-partum estrus in this species. Delayed implantation was proposed based on the observation of a several months lag between copulation and the estimated pregnancy initiation date. The present study demonstrated the advantage of using remote technologies to learn the life history of resting fossorial species.