﻿Magnifying the hotspot: descriptions of nine new species of many-plumed moths (Lepidoptera, Alucitidae), with an identification key to all species known from Cameroon

﻿Abstract This study confirms Mount Cameroon as an unprecedented hotspot for the diversity of many-plumed moths, with the discovery and description of nine new species: Alucitafako Ustjuzhanin & Kovtunovich, sp. nov., Alucitapyrczi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasroczki Ustjuzhanin & Kovtunovich, sp. nov., Alucitapotockyi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasedlaceki Ustjuzhanin & Kovtunovich, sp. nov., Alucitatonda Ustjuzhanin & Kovtunovich, sp. nov., Alucitaerzayi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasokolovi Ustjuzhanin & Kovtunovich, sp. nov., and Alucitahirsuta Ustjuzhanin & Kovtunovich, sp. nov. Additionally, four additional species are reported from the Mount Cameroon area as new for the country: Alucitaagassizi, Alucitadohertyi, Alucitaplumigera, and Alucitarhaptica. Of the 89 Alucitidae known from the Afrotropics, the studied area hosts 36 species, most of which are endemic to the area. This unprecedented level of diversity and endemism within this lepidopteran family highlights Mount Cameroon’s significance as a stronghold for specialised insect taxa. Efficient conservation efforts are necessary to protect these ecosystems and their associated unique microlepidopteran diversity.


Introduction
Mount Cameroon represents a well-documented hotspot of diversity for many-plumed moths (Lepidoptera, Alucitidae), a group of moths distinguished by the division of their wings into six lobes.In our previous studies (Kovtunovich and Ustjuzhanin 2016;Ustjuzhanin et al. 2018aUstjuzhanin et al. , 2020a)), we described 17 new ZooKeys 1193: 25-48 (2024), DOI: 10.3897/zookeys.1193.111544 Peter Ustjuzhanin et al.: Many-plumed moths on Mount Cameroon species of many-plumed moths and reported additional six species within the Mount Cameroon area.By these numbers, Mount Cameroon was revealed as a key hotspot for the group diversity as it hosts a considerable proportion of the 80 species of Alucitidae known from the Afrotropical region prior to this study (De Prins and De Prins 2023).
In this study, we report the remaining material of Alucitidae gathered during our extensive sampling in the Mount Cameroon area between 2014 and 2017 as part of a large ecological project (e.g., Maicher et al. 2020a, b).Nine species of Alucita are described as new for science, and four additional species are reported as new for Cameroon.Furthermore, we furnish an identification key for the majority of species (excluding four species with unknown males) reported from Cameroon.

Materials and methods
Our sampling of Alucitidae was performed in nine rainforest localities situated on the south-western and southern slopes of Mount Cameroon, spanning from November 2014 to October 2017.The sampled elevations ranged from 30 to 2200 m a.s.l.These diverse localities provided a comprehensive range of regionally available rainforest habitats.All reported specimens were attracted to light.A comprehensive sampling protocol was previously outlined in Ustjuzhanin et al. (2018a) and Maicher et al. (2020a).
Holotypes will be housed in the Nature Education Centre, Jagiellonian University, Kraków, Poland (NECJU), while paratypes and other specimens will be divided between NECJU and the personal collections of P. Ustjuzhanin and V. Kovtunovich, located in Novosibirsk and Moscow, Russia (CUK).
For identification, we dissected and examined genitalia of most specimens, adhering to the established protocol described in Ustjuzhanin et al. (2018a).Each permanent preparation received a unique code that allows for convenient retrieval and cross-referencing in the collections where they are stored.The relevant codes are provided in the captions of the genitalia figures.
The sampling localities are listed below in an alphabetic order: Bamboo Camp.Bamboo Camp (350 m a.s.l.), Mount Cameroon (SW slope), 4.0879°N, 9.0505°E; a lowland rainforest with historical disturbances from selective logging.Bimbia-Bonadikombo.Mexico Camp (30 m a.s.l.), Bimbia-Bonadikombo Community Forest, 3.9818°N, 9.2625°E; a littoral forest in the part of the community forest that is officially disturbance-free, but with extensive current logging (Ferenc et al., 2018).Crater Lake.Crater Lake camp (1450 m a.s.l.), Mount Cameroon (SW slope), 4.1443°N, 9.0717°E; a submontane rainforest locally disturbed by forest elephants.Drink Gari.Drink Gari camp (650 m a.s.l.; also known as "Drinking Gari" or "Drink Garri"), Mount Cameroon (SW slope), 4.   Ustjuzhanin & Kovtunovich, 2018 (known so far only from the male, also collected at Bamboo Camp) but can be distinguished by the presence of a pale orange medial band on its hind wings, the orange band on its fore wings being half as wide, and a larger wingspan.In terms of female genitalia, A. pyrczi shares similarities in the structure of antrum and the shape of papillae anales and apophyses with Alucita molliflua (Meyrick, 1927) (Figs 24,25).However, these species are clearly differentiated by the structure of the ductus, the position of the ductus seminalis, and the absence of signa in the bursa copulatrix.

Descriptions of the new species
External characters.The head and thorax are covered with appressed darkgrey scales, while the tegulae appear white.Labial palpi are dark grey on the outside, with white scales on the inner side, and are twice as long as the longitudinal eye diameter.The third segment is short and white, with scattered tiny brown scales.The antenna is yellowish white.The wingspan measures 18 mm.The fore wing is brown with a distinct orange medium band and is lightened with yellowish white scales at the base.The hind wings are noticeably paler than the forewings and feature a pale orange medial band.The lobes of the hind wings have dark-brown and orange spots of scales submarginally.The fringes on the hind wings are whitish apart from sections of dark-brown hairs around the medial band and submarginal spots, and the distal half of the costa of the first lobe of the hindwing is dark brown.The hind legs are pale-yellow.
Female genitalia.The papillae anales are narrow and elongated.The posterior apophyses are shorter than the anterior apophyses.The antrum is wide and goblet-like, sclerotised.The ductus between the ductus seminalis and antrum is narrow and short.The ductus widens significantly at the junction with the ductus seminalis and narrows at its entrance to the bursa copulatrix.The bursa copulatrix is rounded, and no signa are present.
Distribution.The species was found in Cameroon only.Flight period.The species was collected in December.
Etymology.The species is named in honour of the Polish lepidopterist Tomasz Wilhelm Pyrcz, who contributed significantly to the collection and study of butterflies and moths in many parts of the world, including Cameroon.8).However, it can be distinguished from these species by the darkened terminal

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band present on all wings and by its larger size.The most reliable distinguishing feature lies in the male genital structure.While the general structure of the male genitalia is reminiscent of A. seychellensis, A. sroczki has narrower valves, a shorter gnathos, and a set of complex sacculus structures with serrated spiky forms, as well as a narrower aedeagus.In the female genitalia, the new species differs from A. seychellensis and A. megaphimus in the deep V-shaped notch on the outer edge of the antrum.External characters.The head, thorax, and tegulae are covered with clinging grey-brown hairs.Labial palpi are dark-grey and measure 2.5-3 × the longitudinal eye diameter.The third segment is thin, long, and belted basally and apically with narrow white scales.The antennae are dark brown.The wingspan ranges from 16 to 20 mm (holotype 18 mm).All wings exhibit a greyish brown colouration, with four and six distinct pale transverse zigzag bands on the fore wing and hind wing, respectively.The wings are noticeably darkened distally.

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The fringe on the lobes of all wings features alternating portions of pale yellow and dark brown hairs.The hind legs are pale yellow.Male genitalia.The uncus is simple, long, and medially narrowing, with a widened distal end that bears a small notch.The gnathos is slightly shorter than the uncus and narrow, tapering to an acute apex.The gnathos arms are short and wide.The valves are simple and short, measuring half the length of the uncus.The distal portion of the sacculus is expanded as a forked structure.The outer portion of this fork is narrower, internally serrated and terminating in an acute and slightly inwardly bent apex.The inner portion of the fork is wider and finger-like, also serrated on the inside.The basal portion of the sacculus is wide, with a globular sclerotised formation covered with tiny sharp needles.The anellus arms are long, equal in length to the gnathos, and wide at the base, gradually narrowing.The saccus is slightly elongated and caudally rounded.The aedeagus is straight, basally widened, and 1.5 × longer than the uncus.The cornutus is needle-like, distinctive, and occupies most of the aedeagus.
Female genitalia.The papillae anales are narrow and elongated.The posterior apophyses are long and thin, approximately equal in length to the anterior apophyses.The antrum is tubulate and sclerotised, with a narrow V-shaped notch on the outer edge.The ductus is very short, slightly shorter than the antrum, and the ductus seminalis passes from the confluence of the ductus into the bursa.The bursa is oval and very large, featuring two narrow ribbon-like signa.
Distribution.The species was found in Cameroon only.Flight period.The species was sampled in April and October.
Etymology.The species name is a noun in apposition.It is named in honour of the curators from NECJU, Ewelina Sroka and Karolina Sroka, who crucially contributed to the processing of the abundant moth material collected on Mount Cameroon and several other Afrotropical localities.The name 'sroczki' refers to the nickname commonly used for the twin sisters.
Note.Previously, Ustjuzhanin and Kovtunovich (2017) erroneously synonymised A. megaphimus with A. seychellensis.Later, PU and VK re-examined their characters in more detail, and therefore we consider Alucita megaphimus (Hering, 1917), stat.rev., as a separate species.Differential diagnosis.In the male genitalia, the new species exhibits great similarity to Alucita escobari Ustjuzhanin & Kovtunovich, 2018, from which it differs in the more reduced notch in the top of the uncus and in the caudally acute saccus.In contrast, in A. escobari the notch on the top of the uncus is clearly expressed, triangular, and the saccus is caudally smooth, oval, and not acute.Additionally, these two moth species are clearly distinct externally.In the female genitalia, the new species closely resembles Alucita besongi Ustjuzhanin & Kovtunovich, 2018 and Alucita janeceki Ustjuzhanin & Kovtunovich, 2018.From the former, it differs in the oval, elongated bursa copulatrix and the absence of small signa in it, while in A. besongi the bursa copulatrix is pear-like, its surface covered with tiny signa.From the latter, the new species differs in the shape of the bursa copulatrix and the antrum, in the new species the bursa copulatrix narrows caudally, while in A. janeceki, it has a rounded base.The antrum in the new species has a narrow V-shaped notch on the outer edge, while in A. janeceki the notch is wide.From both species, the new species differs in the very long posterior apophyses, in A. besongi and A. janeceki the anterior and posterior apophyses are equal in the length.Furthermore, the male genitalia of the new species are clearly different from those of A. besongi and A. janeceki.

Alucita fako
External characters.The head, thorax, and tegulae are white.Labial palpi are pale-yellow and measure twice the longitudinal eye diameter.The antennae are yellowish white.The wingspan ranges from 9 to 12 mm (holotype 11 mm).The wings are pale yellow, mottled, with alternating white and yellowish brown portions of scales.All lobes of the wings have small dark spots of scales on tips.The fringe on all lobes of the wings has alternating white and pale brown portions of hairs.The hind legs are white.
Male genitalia.The uncus is long, distally extended, and apically with a poorly visible notch.The gnathos is slightly shorter than the uncus and apically acute.The gnathos arms are wide and slightly shorter than the gnathos itself.The valves are wing-like and apically have a bundle of thin needle-like setae.

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The anellus arms are wide, straight, and equal in length to the gnathos.The saccus is elongated and forms a narrow triangle, with an acute tip.The aedeagus is almost straight, obliquely cut apically, and without cornuti.Female genitalia.The papillae anales are narrow and elongated.The posterior apophyses are very long and thin.The antrum is sclerotised, with a narrow V-shaped notch on the outer edge.The ductus is very short, slightly shorter than the antrum, and the ductus seminalis extends distally inside the bursa copulatrix.The bursa copulatrix is oval, elongated, and noticeably narrows at the end, with numerous longitudinal long ribs inside.
Distribution.The species was found in Cameroon.Flight period.The species was collected in May and from October to December.
Etymology.The species is named after Fako, the local name of Mount Cameroon, which is the type locality of the species.The name aims to emphasise the importance of the area and encourage the protection of the species' habitats.Differential diagnosis.In terms of wing colouration, the species somewhat resembles Alucita mischenini Ustjuzhanin & Kovtunovich, 2018, but it differs in the length of the apical dark-brown portion of scales on the first and second lobes of the fore wing.In the new species, the portion on the second lobe is twice as long as that on the first lobe, while in A. mischenini it is equal to or even shorter than that on the first lobe.In the male genitalia, there is also a similarity to A. mischenini, but in the new species, the saccus has a distinct triangular notch caudally, and the aedeagus has an ordered arrangement of needle-like cornuti distally.In contrast, in A. mischenini the notch is absent on the saccus, and the cornuti in the aedeagus are tiny and chaotically disorganised.

Alucita sedlaceki
External characters.The head and thorax are brown, and the tegulae are white.The labial palpi are brown and measure 2.5 × longer than the longitudinal eye diameter.The third segment is short, isolated, and directed upwards.The antennae are yellowish brown and serrated.The wings are white, with black and brown portions of scales.The medial band is well-developed.The wingspan ranges from 12 to 16 mm (holotype 16 mm).The first lobe on the fore wing has alternating brown and yellow rectangular portions.The apical dark brown portion on the first lobe is half as long as the apical portion on the second lobe.The fore wings are basally darkened with dark-brown scales.Medially, they have a wide brown band, which is missing on the sixth lobe.On the hind wings, this band is positioned closer to the base of the wing.All wings have a dark brown subterminal band that is broken in the fifth lobe of the forewing and the third lobe of the hindwing, with the small dark spots of scales subapically on all lobes of all wings.The fringe on the wings is pale, with only the banded portions being brown.The hind legs are yellowish white.
Male genitalia.The uncus is long, basally and medially narrow, and distally widened, with a small notch at the apex.The gnathos is narrow, apically acute, and equal in length to the uncus.The gnathos arms are short and wide.The valves are short, wide, and wing-like.The anellus arms are long, slightly shorter than the gnathos, but significantly wider than it, being basally wide and apically narrowing.The saccus is equal in length to the anellus arms, with a clearly expressed triangular notch caudally.The aedeagus is slightly concave medially and almost equal in length to the entire genital structure (excluding the uncus).The aedeagus large needle-like cornuti distally arranged in an orderly array.
Distribution.The species was found in Cameroon only.Flight period.The species was collected in February, April, and October.
Etymology.The species is named in honour to Ondřej Sedláček, a recognised ornithologist and entomologist with experience from many African countries.On Mount Cameroon, he established several ongoing ecological research projects and was instrumental in helping local communities to understand how to protect the unique local ecosystems in which they live.

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new species stands out with its unusual asymmetric structure of the antrum, which has no analogues among known species.
External characters.The head, thorax, and tegulae are dark brown.The labial palpi are short, straight, and slightly longer than the longitudinal eye diameter.The antennae are brown.The wingspan is 16 mm, and the wings are dark brown.Narrow pale longitudinal bands are present on the lobes of all wings.The fringe on the lobes of all wings is greyish brown.The hind legs are pale yellow.
Female genitalia.The papillae anales are wide.The posterior apophyses are short, thick, and slightly shorter than the anterior apophyses.The antrum is asymmetric, sclerotised, and distally tubulate, with a small triangular notch in the middle.The medium portion of the antrum is very wide, with the right half distinctively protruding to the side, creating a structural asymmetry.The lower portion of the antrum is membranous and bears two round sclerotised plaques.The ductus is short, almost invisible, and passes into a narrow membranous bursa copulatrix, with no signa observed.
Distribution.The species was found in Cameroon only.Flight period.The species was collected in February.
Etymology.The species name is a noun in apposition, given in honour of Antonín "Tonda" Tropek, who is RT's father.longipenis, the aedeagus of the new species is very long, but it is slightly shorter, has fewer curves, and is straight for the distal portion.Additionally, the saccus of A. erzayi is gently curved distally and apically acute, while in A. longipenis the saccus is bent and apically acute.The wingspan of the new species is 14 mm, whereas it is 18-23 mm for A. longipenis.

Alucita erzayi
External characters.The head, thorax, and tegulae are yellowish white.The labial palpi are thin, straight, and twice as long as the longitudinal eye diameter.The antennae are pale yellow.The wingspan is 14 mm, and the wings are pale yellow, interspersed with brown strokes and spots.Two small dark brown patches are present in the basal portion of the costa of the first lobe of the fore wing.Indistinct pale brown regions of scales are present in the medial and distal portions of the first two lobes.The fringe on the wings is pale yellow, and the hind legs are pale yellow.
Male genitalia.The uncus is relatively long and evenly wide throughout its length, with a rounded apex.The gnathos is long and narrow.The valves are reduced.The anellus arms are long and evenly narrow throughout their length.The saccus is long, elongated, and smoothly bent caudally, with a clearly acute apex.The aedeagus is very long, ~ 4 × longer than the entire genital structure, forming two arched curves in the medium part.No cornuti are present.
Distribution.The species was found in Cameroon only.Flight period.The species was collected in April.
Etymology.The species is a noun in apposition.'Erzayi' is a word in the Bakweri language, which is the dominant local language in the Mount Cameroon region, and it translates to "feather".This corresponds with the appearance of many-plumed moths' feather-like characteristic wing lobes.

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apically framed with a white subapical zigzag.The hind wings are noticeably paler than the fore wings, with a brown band widening towards the last three lobes.There are bundles of brown hairs on the lobes, both distally and basally, with white fringes between them.The hind legs are pale yellow.Male genitalia.The uncus is long, noticeably exceeding the gnathos in length, and is distally slightly widened with a small apical notch.The gnathos is narrow and apically acute.The valves are long and membranous, apically smoothly forming a wide oval shape.The anellus arms are thin and straight.The saccus is caudally oval.The aedeagus is short, almost straight, and has two spiky cornuti.
Distribution.The species was found in Cameroon only.
Flight period.The species was collected in February, April, and May.
Etymology.The new species is named after Vasily Igorevich Sokolov (Moscow, Russia), the famous Russian ichthyologist and bioresource recovery specialist.Differential diagnosis.The mushroom-like antrum and elongated crest-like signum of this species bear similarity to Alucita ectomesa (Hering, 1917) (illustrated in Ustjuzhanin and Kovtunovich 2016), but it can be distinguished by the wider ductus, the round bursa copulatrix, and numerous tiny spiky signa present in it.Additionally, the unique colouration of the wings sets A. hirsuta apart from all other African Alucitidae species.

Alucita hirsuta
External characters.The head, thorax, and tegulae are dark brown.The labial palpi are short, slightly longer than the longitudinal eye diameter.The antennae are brown.The wingspan is 14 mm, and the wings have a reddish brown appearance.The lobes of all wings bear protruding tousled dark-brown hairs, especially dense on the first two lobes of the fore wings, creating the appearance of a shaggy moth.There are narrow, poorly visible pale longitudinal bands on all wings.The fringe on all wings ranges from pale to dark brown.The hind legs are yellow.
Female genitalia.The papillae anales are narrowly triangular in shape.Both the posterior and anterior apophyses are of equal length, thick, and straight.The antrum is wide and mushroom-like.The ductus is wide, corrugated, and strewn with narrow strands.The ductus seminalis passes from the middle of the ductus.The bursa copulatrix is round, with a robust crest-like signum located in the upper part of the bursa, near the confluence of the ductus.Numerous tiny spiky signa densely cover the entire surface of the bursa.
Distribution.The species was found in Cameroon only.Flight period.The species was collected in October.
Etymology.The species name is derived from Latin 'hirsute' (shaggy, bristly, hairy).It refers to the appearance of the adult moth, with tousled protruding dark-brown hairs on the wings, reminiscent of a hairy and shaggy moth.Differential diagnosis.The yellow wing colour of this species resembles Alucita compsoxantha Meyrick, 1924, but it can be distinguished by the differences in the position of the bands.In the male genitalia, A. potockyi shares similarities with Alucita tesserata (Meyrick, 1918) in the short uncus widening towards the apex and in the saccus shape, but it clearly differs in the apically tapered gnathos, long narrow valves and the long aedeagus that exceeds the genital structure in its length.In comparison, A. tesserata has a gnathos that strongly widens apically, short and wide valves, and a noticeably smaller aedeagus relative to the genital structure (see Ustjuzhanin et al. 2020a for the genitalia illustration).

Key to identify Alucita species in the Mount Cameroon area (males only)
In addition to the 36 species of Alucita previously reported from the Mount Cameroon area in Ustjuzhanin et al. (2018aUstjuzhanin et al. ( , 2020a)), Ustjuzhanin and Kovtunovich (2016), and in this study, the identification key also includes the only other species known from Cameroon, Alucita illuminatrix (Meyrick, 1929) described from Bitje, South Region, Cameroon (DePrins and DePrins 2023).Although this species has not been recorded in the Mount Cameroon area, we have included it in the identification key.On the other hand, Alucita tatjana Ustjuzhanin & Kovtunovich, 2020, A. pyrczi, A. tonda, and A. hirsuta

Discussion
The Mount Cameroon area hosts a remarkable diversity of Alucitidae.This paper adds nine newly described species and four other newly reported species, bringing the total count to 36 species.This comprises 40% of all 89 Alucitidae species known from the Afrotropical region, including 80 species listed in the Afromoths database (De Prins and De Prins 2023) and nine species described in this study.The extent of local diversity in this moth group is unprecedented, as only a few species of this group are known from any other locality in the region (Ustjuzhanin et al. 2018a(Ustjuzhanin et al. , 2020a;;De Prins and De Prins 2023).Mount Cameroon is known to harbour high diversity in many taxa, including other Lepidoptera groups (e.g., Ballesteros-Mejia et al. 2013;Maicher et al. 2016;Przybyłowicz et al. 2019;Delabye et al. 2020, Mertens et al. 2021).The region's exceptional species richness is often attributed to its location at the confluence of the Guinean and Congolian biogeographic regions, and a presence of diverse habitats along its elevational and precipitation gradients (Cable and Cheek 1998;Bergl et al. 2007;Hořák et al. 2019;Maicher et al. 2020a;Delabye et al. 2021;Doležal et al. 2022).Additionally, the area's relatively high isolation further contributes to the unique ecological conditions (Ustjuzhanin et al. 2018a).However, the sheer magnitude of many-plumed moth species richness on Mount Cameroon surpasses expectations based on combinations of these exceptional factors in other Afrotropical localities.Despite this unique combination of conditions, it remains challenging to fully account for why Mount Cameroon exhibits such a substantial predominance of many-plumed moths compared to all other known sites in the Afrotropical region.
Notably, 24 of these species have been described solely from the Mount Cameroon area (as A. ludmila was already known from Nigeria and Ghana when it was described; Ustjuzhanin et al. 2018a), and the majority of them (except for A. mischenini and A. zinovievi recently reported from Liberia and Ghana, respectively; Ustjuzhanin et al. 2020b) are considered endemic to Cameroon and have not been recorded elsewhere.This level of endemism among the Alucitidae underscores the importance of Mount Cameroon as a vital refuge for specialised and unique insect taxa.While the region is already renowned for its rich endemic diversity of moths and butterflies (e.g., Larsen 2005;Sáfián and Tropek 2016;Przybyłowicz et al. 2019;Sáfián et al. 2019), as well as its endemic plant (Cable and Cheek 1998) and vertebrate (Fjeldså and Lovett 1997;Bergl et al. 2007) species, the unparalleled level of endemism of Alucitidae further highlights the area's significance as a centre of microlepidopteran diversity in the Afrotropics.
The discovery of such a diverse and endemic locality on Mount Cameroon has exceeded expectations, despite the limited knowledge of Afrotropical microlepidoptera.The implications of this exceptional diversity extend beyond taxonomy, prompting future research into the evolutionary and ecological processes that have facilitated the development of this diverse moth community.The unique diversity and endemism of many-plumed moths in the Mount Cameroon area underscore the urgent need for efficient conservation of ecosystems and habitats in the region, especially considering previous conservation efforts that faced challenges in some parts (Ferenc et al. 2018).Collaborative efforts among researchers, conservationists, and local communities are crucial in preserving this treasure trove of biodiversity and securing the future not only for many-plumed moths.
Elephant Camp (1850 m a.s.l.), Mount Cameroon (SW slope), 4.1170°N, 9.0729°E; a montane forest with a sparse canopy layer as a consequence of natural disturbances by forest elephants.Mann's Spring.Mann's Spring camp (2200 m a.s.l.), Mount Cameroon (SW slope), 4.1428°N, 9.1225°E; a montane forest at the natural timberline.