Research Article |
Corresponding author: Rob Felix ( robfelix1@gmail.com ) Academic editor: Axel Hochkirch
© 2025 Rob Felix, Annelies Jacobs, Michel Lecoq.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Felix R, Jacobs A, Lecoq M (2025) Rediscovery of the Monte Gordo Grasshopper Eyprepocprifas insularis: An ancient brachypterous species endemic to São Nicolau, Cape Verde (Orthoptera, Eyprepocnemidinae). Journal of Orthoptera Research 34(2): 159-168. https://doi.org/10.3897/jor.34.144016
|
The Monte Gordo Grasshopper Eyprepocprifas insularis, considered extinct, was rediscovered on the island of São Nicolau, Cape Verde, in 2023. This unique brachypterous grasshopper is a “living fossil” endemic to the archipelago. This paper provides updated taxonomic information, describes the female for the first time, examines habitat characteristics and the impact of human activity, and proposes an IUCN Red List status. The study emphasizes the need for further research on this resilient yet threatened species.
Africa, extinct, grasshopper, highlands, Red List, taxonomy
Eyprepocprifas insularis Donskoff, 1982 was first discovered by ML and colleagues of the Programme de Recherche Interdisciplinaire Francais sur les Acridiens du Sahel (PRIFAS) in October 1980 on the island of São Nicolau, Cape Verde. It is the only brachypterous grasshopper endemic to this archipelago. Only one male specimen was found, and despite long periods spent on the islands between 1979 and 1982 by ML, no other specimen was encountered. Therefore,
During a trip to São Nicolau from 23–28 Jan. 2023, RF and AJ rediscovered the species and found it present at several sites in Monte Gordo Natural Park (Fig.
Topography.—The Cape Verde Archipelago is situated in the Atlantic Ocean, approximately 600 km west of mainland Africa. It consists of ten islands and several smaller islets formed through volcanic activity associated with a stationary mantle plume beneath the Cape Verde Rise, an elevated region of the ocean floor of volcanic origin (
Cape Verde belongs to the Macaronesian ecoregion, together with Madeira, the Azores, the Canary Islands, and the Savage Islands. The most eastern islands (Sal, Boa Vista, and Maio) are geologically older than the western islands and are more eroded, flat, and topographically monotonous. The western islands (Santo Antão, São Vicente, São Nicolau, Santiago, Fogo, and Brava) have steep, high mountains and a rough topography with deep river valleys or ribeiras (
São Nicolau is located between the islands of Razo and Sal in the northern part of the archipelago (Fig.
Climate.—The climate on São Nicolao is mainly influenced by the cold Canary current and the northeastern trade wind, less so by a southwestern monsoon. The temperature is balanced by the surrounding ocean and ranges from an average of 22°C during the day in February to 27°C in September. In high mountains, the climate is less influenced by the ocean, and frost may occur in winter. The monsoon blows from August until October and is the main cause of the rainy season, which is irregular and limited. Annual precipitation is marked by low and irregular levels of rainfall ranging from 80–300 mm (lowlands) to more than 1,600 mm (highlands). Rain may be absent for several years, causing catastrophic droughts. The northeastern trade wind carries medium humidity, particularly in autumn and winter, mainly affecting mountains above 600 m.a.s.l. This results in fog, which is of utmost importance for additional water supply to the natural vegetation (
Habitats.—Large areas in Cape Verde are covered by open grassland and semi-desert vegetation, and considerable parts of the eastern islands are desertified. Monte Gordo Natural Park on São Nicolau comprises an important humid mountain ecosystem. Significant dense scrublands are characterized by the endemic Euphorbia tuckeyana and other endemic taxa such as Dracaena draco caboverdeana, Asteriscus smithii, and Sideroxylon marginatum. The presence of these species is thought to reflect the composition of local vegetation before human colonization.
Faunistic taxa endemic to São Nicolao and occurring in Monte Gordo are the following reptiles: Hemidactylus nicolauensis Vasconcelos, Köhler, Geniez, & Crochet, 2020, Tarentola nicolauensis Schleich, 1984, and Chioninia nicolauensis (Schleich, 1987) (
The human impact on natural habitats has been considerable for over 500 years, and most vegetation is severely disturbed. The combination of harsh climate and human disturbance limits the regeneration potential of the vegetation, and only remnants of supposedly natural vegetation are left. Monte Gordo Natural Park holds an extensive afforested area planted in the 20th century, mainly comprising the introduced Pinus, Eucalyptus, and Cupressus (
Identification and taxonomy.—The specimens were identified using the original species description published by
Institutional abbreviations.—
RFPC Rob Felix Private Collection, Nijmegen, the Netherlands.
Terminology.—Terminology of the morphological characters follows
Red List assessment.—The area of occupancy (AOO) of Eyprepocprifas insularis was calculated by summing the total area of occupied 2×2 km grid squares. The lower extent of occurrence (EOO) was determined using the minimum convex hull around the confirmed occurrences, while the upper EOO was calculated using the minimum convex hull encompassing the entire estimated geographical range, which may overestimate its actual distribution.
The estimated geographical range of E. insularis was derived from our field records, habitat information gathered during our survey, a digital elevation model (
Due to data limitations, applying all five IUCN criteria categories was impossible. Specifically, there is no information on population status, trends, or sufficient data for quantitative analyses. Therefore, the assessment is limited to criteria B and D2, which rely on the geographic distribution parameters AOO and EOO. Based on these values and information on threats and habitat characteristics entered into the Species Information Service (SIS), IUCN’s central database, the appropriate threat category was automatically determined.
Acrididae
Eyprepocnemidinae
Eyprepocnemidini
Type material. Holotype. CAPE VERDE • 1♂; São Nicolau; [670 m.a.s.l.]; [16.6182°N, 24.3324°W]; Oct. 1980; PRIFAS exp. leg.;
Additional material. CAPE VERDE • 1♂; São Nicolau, Cachaço, Monte Cintinha; 670 m.a.s.l.; 16.6246°N, 24.3290°W; 23 Jan. 2023; R. Felix leg.; RFPC • 1♂, 1♀ nymph; São Nicolau, Cachaço, Monte Cintinha; 700 m.a.s.l.; 16.6243°N, 24.3297°W; 23 Jan. 2023; R. Felix leg.; RFPC • 1♂; same data as for previous; RFPC RMNH.INS1622466 • 1♂, 1♀; São Nicolau, Cachaço, Monte Cintinha; 670 m.a.s.l.; 16.6246°N, 24.3290°W; 24 Jan. 2023; R. Felix leg.; RFPC • 1♀; São Nicolau, Cachaço, Monte Cintinha; 710 m.a.s.l.; 16.6236°N, 24.3307°W; 24 Jan. 2023; R. Felix leg.; RFPC RMNH.INS1622467 • 1♂; São Nicolau, Road Tarrafal – Cachaço, Near pass; 800 m.a.s.l.; 16.6182°N, 24.3324°W; 26 Jan. 2023; R. Felix leg.; RFPC.
Eyprepocprifas insularis is relatively robust and rough and has a pronotum with a distinctly raised median carina; otherwise, it is a typical member of the Eyprepocnemidinae Brunner von Wattenwyl, 1893. Members of this subfamily are characterized by a subconical head with a triangular profile, the absence of the external (pre-)apical spine of the hind tibiae, and the specific morphology of the male internal genitalia. Most eyprepocnemidine genera have bilaterally compressed male cerci with a downwardly curved apex (
To some extent, female Eyprepocprifas resemble the nymphs of Heteracris littoralis (Rambur, 1838), a species common in Cape Verde at lower altitudes. Female Eyprepocprifas have longer antennae, a longer ovipositor, a less globose head, and a median carina of the pronotum that is much more raised. The characters mentioned earlier in this paper easily distinguish it from any other Cape Verdean acridid congener.
Morphometrics of Eyprepocprifas insularis
Parameters | Male (N = 6, including holotype) | Female (N = 2) |
---|---|---|
Body Length (frons–subgenital plate) | 19.0 (18.2–19.5) | 31.1 (28.8–33.4) |
Length Hind Femur | 13.3 (12.8–13.9) | 20.8 (20.7–20.8) |
Length Pronotum | 4.6 (4.2–5.0) | 8.7 (8.5–8.8) |
Length Prozona | 2.7 (2.5–2.8) | 4.6 (4.5–4.7) |
Length Metazona | 2.0 (1.7–2.2) | 4.1 (4.0–4.1) |
Eye Height | 2.5 (2.4–2.7) | 3.2 (3.0–3.3) |
Eye Width | 1.4 (1.3–1.4) | 1.7 (1.6–1.8) |
Interocular Space | 0.7 (0.7–0.8) | 1.3 (1.2–1.3) |
Length Tegmen | 5.1 (4.6–5.9) | 9.5 (9.1–9.8) |
Integument granular, except for smooth ventral surface. Head subconical and triangular in profile, inclined at a 60° angle to the horizontal. Face narrow (2 mm) and elongated (3.5 mm). Fastigium triangular, slightly broader than long, rounded at front, and flat. Frontal ridge highly prominent, measuring 1 mm at fastigium, its base narrow (0.8 mm), and tapering upward, with minimal constriction at the median ocellus. Temporal foveae absent. Lateral ocelli small, positioned beneath the fastigium margin, invisible from above. Interocular space equals width of antennal scape (0.7 mm). Antennae filiform, composed of 22 segments, reaching halfway the tegmina. Eyes oval (2.4 mm/1.3 mm), positioned 1 mm above the genal suture.
Pronotum tectiform, with a raised and horizontal median carina. Lateral carinae well-marked, diverging posteriorly. Three deep transverse sulci cut the median keel. Prozona (2.6 mm) longer than metazona (1.9 mm). Hind margin of the pronotum angular, rounded at the apex, barely notched. Lateral lobes trapezoidal, as high as ventral width (2.8 mm). Prosternal tubercle large, conical, narrow at the base, rounded at the apex. Mesosternal space subquadrate; mesosternal lobes rounded.
Tegmina short but variable in length, extending beyond the hind margin of the second or third abdominal tergite; narrow, 2 mm in width; nearly in contact along dorsal edges, tapering anteriorly and posteriorly from the midpoint, apex angularly rounded. Hind wings present, reaching the first abdominal tergite’s hind margin.
Hind femora elongated, 4.5 times longer than wide, exceeding the abdominal extremity. Upper basal lobe larger and longer than lower one. Lower genicular lobes acute; Brunner’s organ well developed. Fore and mid tibiae ventrally with one external apical spine in addition to the six spines on the external margin and five and seven spines on the internal margin, respectively.
Hind tibiae with nine spines on the inner and 9–10 spines on the outer dorsal margins. Internal spines and spurs twice the length of the external ones. Tibiae shorter than the femora; arolium of all tarsi large, slightly longer than half the length of the claws. Last segment of fore and mid tarsi as long as the other two combined; last segment of the hind tarsi as long as first segment alone.
Abdominal tympanum large and well-exposed. Abdominal tergites 1–7 with a raised median ridge. Tergite X incurved with a shallow furcula. Epiproct triangular, slightly elongated, rounded at the apex, fairly flat, with two short, low, flat longitudinal ridges near the base on either side of the midline; twice as long as wide. Paraproct with a short anterior projection (
Male: Very variable in color and design: Pronotal disc varies from nearly plain beige with some dark markings (Fig.
Epiphallus consists of two small sub-square lateral plates connected by a short, slightly arched bridge. Ancorae large, articulated on the lateral plates, sharp, and prominently projecting. Lophi large and bilobed, with hind lobe broad, horizontal, slightly bilobed, and vaulted. Internal lobe of lophi globular and spiculated. Oval sclerites small and triangularly rounded; lateral sclerites long, narrow, and triangular, tips pointing downward (Fig.
Eyprepocprifas insularis Donskoff, 1982, female and male phallic complex. A Female; B. Ovipositor dorsal view; C. Ovipositor lateral view; D. Male phallic complex dorsal view (anterior end on top); E. Male phallic complex lateral view (anterior end on the left). An: Ancorae, Br: Bridge, Lo: Lophi. Monte Cintinha, São Nicolau, Cape Verde, Jan. 2023. Female: RMNH.INS1622467; Male: RMNH.INS1622466. Scale bars: 1 cm (A); 1 mm (B–E). Photo credit: Luc Willemse, NBC, Leiden.
Female: Same characters as male, with the following differences: Total length 28.8–33.4 mm; 1.5–2.0 times larger than male (Table
Eyprepocprifas insularis is endemic to the island of São Nicolau, Cape Verde. After its discovery in 1982, it was considered extinct by
In January 2023, it was found at several localities in Monte Gordo Natural Park (Fig.
Sightings of Eyprepocprifas insularis Donskoff, 1982, São Nicolau, Cape Verde, January 2023. All records, including the collected specimens mentioned in the Material section, are shown. The location numbers refer to those in Fig.
Date | Location | Elev. m.a.s.l. | N | Sex | Stage | Lat, Long E |
---|---|---|---|---|---|---|
23-1 | Monte Cintinha (Loc. 2) | 670 | 1 | ♂ | Adult | 16.6246°N, 24.3290°W |
23-1 | Monte Cintinha (Loc. 2) | 700 | 3 | ♂ | Adult | 16.6243°N, 24.3297°W |
23-1 | Monte Cintinha (Loc. 2) | 700 | 1 | ♀ | Nymph | 16.6243°N, 24.3297°W |
24-1 | Monte Cintinha (Loc. 3) | 660 | 1 | ♂ | Adult | 16.6262°N, 24.3267°W |
24-1 | Monte Cintinha (Loc. 2) | 670 | 1 | ♀ | Adult | 16.6246°N, 24.3291°W |
24-1 | Monte Cintinha (Loc. 2) | 670 | 1 | ♂ | Adult | 16.6246°N, 24.3291°W |
24-1 | Monte Cintinha (Loc. 2) | 710 | 1 | ♀ | Adult | 16.6236°N, 24.3307°W |
25-1 | Assomada Pass (Loc. 4) | 650 | 2 | ♂ | Adult | 16.6397°N, 24.3712°W |
25-1 | Assomada Pass (Loc. 4) | 650 | 1 | ♂ | Adult | 16.6398°N, 24.3705°W |
25-1 | Assomada Pass (Loc. 4) | 650 | 1 | ♂ | Adult | 16.6397°N, 24.3707°W |
26-1 | Road Tarrafal–Cachaço, near pass (Loc. 1) | 800 | 1 | ♂ | Adult | 16.6182°N, 24.3324°W |
27-1 | Monte Gordo (Loc. 6) | 1100 | 1 | ♂ | Adult | 16.6213°N, 24.3490°W |
27-1 | Monte Gordo (Loc. 5) | 1050 | 4 | ♀ | Nymph | 16.6279°N, 24.3543°W |
Eyprepocprifas insularis is a montane species restricted to the higher parts of the island of São Nicolao. All records are from elevations between 650–1100 m.a.s.l. Table
Site descriptions with sightings of Eyprepocprifas insularis Donskoff, 1982 on São Nicolau, Cape Verde in January 2023. Numbers correspond with those in Fig.
1. | Road from Tarrafal–Cachaço, near the pass 800 m.a.s.l. Steep, rocky northeastern slope with scrub vegetation containing Euphorbia tuckeyana and Furcraea foetida. Mostly dry, with some wetter gullies (Fig. |
2. | Trail eastwards from Capela de Nossa Senhora do Monte Cintinha 650–750 m.a.s.l.: dry, warm, southeastern slope with open, scrubby, and grassy vegetation alternated with more densely vegetated patches with scrubs and Arundo donax (Figs |
3. | Part of the trail eastwards from Capela de Nossa Senhora do Monte Cintinha, where it bends to the northeastern slope 660 m.a.s.l. The cooler and humid conditions make it more densely vegetated with higher scrubs of Euphorbia tuckeyana. |
4. | Assomada Pass, between 650 m.a.s.l. Steep, northeastern slope with Euphorbia tuckeyana scrubs (Fig. |
5. | The northeastern slope of a deep valley with densely woody vegetation at 1050 m.a.s.l. with Pinus, Eucalyptus, and undergrowth of Euphorbia tuckeyana. Nymphs were found here on Asteriscus smithii Figs |
6. | Steep northeastern slope at 1100 m.a.s.l. with Pinus, Eucalyptus, and Cupressus with undergrowth of Euphorbia tuckeyana. |
Most records are from northeastern, southeastern, and north-facing slopes in the open to more densely vegetated habitats, varying from relatively dry to moist soils. All records are from sites where the native Euphorbia tuckeyana is present; however, several sites are dominated by introduced taxa such as Pinus, Eucalyptus, Cupressus, and Arundo donax (Fig.
All sightings of adults were done on the ground; when flushed, they sometimes landed in the vegetation. The species at least appears to be active at night, but probably also during daytime. The first specimens were found at night, sitting on the ground and rocks, at a site visited before during the day without any sightings of the species. Later, specimens were also found during the day, but maybe these were disturbed before their discovery. Small nymphs were found on Asteriscus smithii (Figs
The holotype was found in October. In January 2023, adults and larger and smaller nymphs were found (Fig.
Habitats of Eyprepocprifas insularis Donskoff, 1982 on São Nicolau, Cape Verde. A. Type locality (locality 1); B. Trail at Monte Cintinha (locality 2); C. Trail at Assomada Pass with Euphorbia tuckeyana (locality 4); D. Patch of forest with Pinus, Eucalyptus and Cupressus in Monte Gordo NP (locality 6). Photo credit: Rob Felix.
Records of Eyprepocprifas insularis Donskoff, 1982 on São Nicolau, Cape Verde. New records are indicated with red dots, and the type locality is indicated with a yellow star. Numbers refer to site descriptions in the text. Monte Gordo Natural Park is indicated in dark green. Base map created using Stamen Design in QGIS.
Accompanying species in the same habitat were Diabolocatantops axillaris (Thunberg, 1815), Cycloptiloides canariensis (Bolívar, 1914), two species of Oecanthus, Phaneroptera sparsa Stål, 1857, Ruspolia fuscopuncata (Karny, 1907), Acanthogryllus sp. and Gryllus bimaculatus De Geer, 1773.
The AOO, calculated from the known data (ca. six sites), comprises 12 km2. The EOO, calculated from the known data, is 4 km2 (Lower EOO), while the maximal estimation is 35 km2 (Upper EOO). The area of the polygon with the estimated geographical distribution (possibly extant) in Fig.
Cape Verdian Orthoptera.—Apart from Eyprepocprifas Donskoff, 1982 from São Nicolao, several other Orthoptera taxa are endemic to Cape Verde: Sphingonotus atlanticus (Popov, 1984) is endemic to the island of Santa Luzia, while several subspecies of wider-distributed species are endemic to various islands.
According to the most recent overview, 42 species of Orthoptera are known from the Cape Verde Archipelago: 28 species of the Caelifera suborder (
Living fossil.—The rediscovery of the only endemic brachypterous grasshopper, Eyprepocprifas insularis, on São Nicolau, an island with a volcanic origin dating back approximately five million years, provides significant insight into the island’s ecological and evolutionary history.
This species, first described as the only unequivocal case of absolute endemism in Cape Verde, is morphologically distinct and not closely related to known African counterparts, suggesting a long evolutionary isolation (
Its limited dispersal capability, attributed to brachypterism, supports the hypothesis that it colonized the archipelago during glacial periods with lower sea levels and reduced distances between the islands and the continent (
While it has been shown that the grasshopper fauna of Cape Verde is probably in perpetual evolution under the combined influence of periods of intense drought, which can eliminate the populations of certain species on one or more islands, and prevailing easterly winds, which can bring new populations/species from the neighboring continent (
Mounting specimens of E. insularis proved quite challenging due to the remarkable difficulty of piercing them with a pin. Surprisingly, an exceptional amount of force was required because the integument appeared extraordinarily thick. This might suggest an adaptation to arid conditions, where a reinforced exoskeleton could provide advantages such as reduced water loss or enhanced resilience to environmental stress. This phenomenon exists in other insects, such as Solenopsis Westwood, 1840 ant species in the southeastern USA (Hymenoptera: Formicidae) (
Threats.—The highland vegetation, the native habitat of E. insularis, has undergone significant changes over time. Before human settlement (~5900–410 cal yr BP), the highlands were dominated by native woody species such as Euphorbia tuckeyana and Dracaena draco subsp. caboverdeana. Pollen studies show that this scrubland was relatively stable and supported a high diversity of endemic herbaceous and sub-scrub plant species, epiphytic ferns, and a variety of fungal communities, forming a humid wooded landscape (
It remains unclear which aspects of the life cycle of E. insularis are affected by habitat alterations and which aspects represent key factors herein. For instance, the decline of native host plants or essential fungi as food sources may be driving factors. Increasing research suggests that fungi are a significant food source in invertebrates (
Several conservation actions have been promoted and implemented, such as reforestation with native plants, removing invasive plant species, and grazing control to protect native vegetation (
Despite the above-mentioned challenges, E. insularis demonstrates resilience, surviving in degraded habitats dominated by introduced plant species. Furthermore, a significant part of its habitat is protected as a natural park (Fig.
Future study.—The rediscovery of this species is a crucial first step toward its conservation. Further research is needed to determine the population size, exact range, ecology, and bottlenecks related to threats to its habitat, such as climate change.
In addition, during our short visit, several new Orthoptera taxa to Cape Verde were recorded that are not yet listed in the above-mentioned references: Oecanthus dulcisonans Gorochov, 1993 (Oecanthidae) on Santiago and Cycloptiloides canariensis (Bolívar, 1914) (Mogoplistidae) and Ruspolia fuscopuncata (Karny, 1907) (Tettigoniidae) on São Nicolau. The discovery of these species underscores the potential for future research in the archipelago.
Comparative studies, particularly in high-altitude and vegetative settings like Monte Gordo Natural Park on other islands like Santo Antão, may uncover related taxa or even undiscovered populations of Eyprepocprifas.
The authors declare that no competing interests exist.
Rob Felix: conceptualization, formal analysis (Red List), investigation, resources, data curation, writing – original draft, visualization; Annelies Jacobs: investigation, writing – original draft, writing – review and editing; Michel Lecoq: writing – original draft, writing – review and editing. All authors have read and agreed to the published version of the manuscript.
We thank Christophe Hervé and Laure Desutter-Grandcolas (Muséum national d’Histoire naturelle, Paris) for providing photographic material of the holotype of E. insularis in