ABSTRACT
Dry evergreen Afromontane forests (DAF) once covered most of the Ethiopian highlands. Currently, they are found as a few patches. DAF restoration is a national forest restoration priority in Ethiopia. It has also been identified to be among global ecosystem restoration priorities. There are sufficient data to show DAF restoration from the soil seed bank or seed rain is hardly possible. Planting trees from multiple provenances is important to restore climate-resilient DAF. Therefore, DAF restoration cannot be achieved through natural regeneration alone and must be accomplished by planting characteristic DAF tree species. The poor soil moisture and nutrient conditions in Ethiopia’s dry highlands result in low native tree seedling survival and growth. Hence, proper preparation of the planting sites and after-planting seedling care for at least two years is a reliable mechanism for achieving successful survival and growth. Along with seedling care after planting, four additional mechanisms discussed in this review (i.e., the use of nurse trees/shrubs, design of nursery practices, inoculation with arbuscular mycorrhizal fungi, and application of biochar) have also been shown to be useful. However, they cannot substitute for proper planting site preparation and post-planting seedling care. The complementary use of these five mechanisms could further enhance success. However, there are more questions than answers regarding the effectiveness of the five mechanisms discussed. We hope this review will motivate researchers to engage further to answer some of the important questions regarding restoration success.
- arbuscular mycorrhizal fungi (AMF)
- assisted natural regeneration (ANR)
- nurse trees
- protected areas
- tropical dry forests
Since the Rio Summit in 1992, biodiversity has emerged as a major global agenda. In 2019, the United Nations General Assembly declared 2021–2030 to be the Decade on Ecosystem Restoration (United Nations 2019). Restoring its dry evergreen Afromontane forests (DAF) is perhaps the most important contribution Ethiopia could make to the global restoration effort. Ethiopia possesses, by far, the largest area suitable for DAF restoration in Africa (Yalden 1983, Bussam 2006). In the past, this area was predominantly covered by DAF, but currently only a few of the original forests remain (Teketay and Anders 1995, Friis et al. 2010). By replanting lost and degraded DAF, Ethiopia could restore a significant proportion of the world’s tropical dry forests (Lemenih and Itanna 2004) and could contribute significantly to global biodiversity conservation and climate change mitigation (Strassburg et al. 2020). Restoring the degraded and lost DAF is also among Ethiopia’s national biodiversity priorities (NBSAP 2005). Whereas Ethiopia has committed to restoring forest on 22 million ha of degraded lands by 2030 (MEFCC 2018), DAF restoration, in particular, has been identified as contributing substantially towards the success of this commitment (MEFCC 2018, Pedercini et al. 2021).
In Ethiopia, the practice of DAF restoration goes back to the 15th century when the Menagesh-Suba Forest was restored by wildlings (i.e., naturally regenerating seedlings) from the Wof-Washa Forest (Sertse et al. 2011). In the 1890s (Bekele 2003) and the 1970–80s (Bishaw 2001), there were massive and successful re-greening and community forestry programs, although these endeavors mainly involved planting exotic trees, not native vegetation.
In the past 30 years, mainly since 2000, DAF restoration has centered on exclosures. Exclosures protect restoration sites from livestock and human interference to facilitate regeneration of natural vegetation. Revegetation potentially can be achieved either passively by allowing natural vegetation to recover, or actively through the construction of soil and water conservation structures and supplemental tree planting (Lemenih and Kassa 2014, Haile and Gebregziabher 2020).
Exclosures have resulted in significant positive environmental and socio-economic impacts in Ethiopia (Lemenih and Kassa 2014, Haile and Gebregziabher 2020). However, the Ethiopian dry highlands are nutrient- and moisture-stressed, the soil lacks seeds of native tree species, and recruitment from seed rain is unlikely (Lemenih and Teketay 2004, Lemenih et al. 2005, Aerts et al. 2007, Asmelash et al. 2021c). While theoretically feasible, passive DAF restoration realistically is not possible. When exclosures have been augmented by tree planting, limited species selection (Reubens et al. 2011) and poor seedling survival (Bishaw 2001, Asmelash et al. 2019) have significantly hampered success. Therefore, devising mechanisms for enhancing the survival and growth of characteristic DAF trees is crucial.
DAF restoration programs would be further enhanced through effective management of remnant DAF patches (Lemenih and Teketay 2004). Therefore, before discussing some of the restoration mechanisms that have been tested for establishing tree species characteristic of DAF, we evaluate the conservation status of the remnant DAF. We also review and discuss important forest restoration concepts relevant to DAF restoration. Our objective is to highlight some of the challenges related to DAF restoration and propose a strategy for effective restoration of the plant community.
Methods
We first made a list of DAF occurring in Ethiopia based on an Ethiopian Biodiversity Institute publication (NBSAP 2005), and then conducted Google Scholar searches using the listed DAF names to identify articles pertinent to the Ethiopian DAF patches. Secondly, we downloaded those articles located using the DAF names, and we also downloaded some of the important references cited in those articles. In so doing, we expanded the DAF list and gathered information relevant to DAF restoration. We used the terms: assisted natural regeneration (ANR), dry evergreen Afromontane forests, ecological restoration, ecosystem restoration, nurse trees, priority effect, protected areas, seedlings regeneration, soil seed bank, tropical dry forests, tropical montane forest, vegetation succession, and Ethiopia, to identify most of the references cited in this review by searching Google Scholar and the Society for Ecological Restoration (SER) databases. No exclusion criteria were set; if relevant information regarding DAF restoration was obtained, every article had equal probability of inclusion. When two or more articles provided similar information, those articles published in the most scholarly journals were given priority for citation.
Conservation Status of Ethiopian DAF
Ethiopia’s dry evergreen Afromontane forests are found in the dry highlands (as indicated by the aridity index; Lemenih and Bongers 2011). They are an integral part of the dry evergreen Afromontane forest and grassland complex ecoregion (DAF-G). Floristically, the DAF-G is the second-richest of the ecoregions in Ethiopia (Friis et al. 2010) and is recognized as one of 35 global biodiversity hotspots (Mittermeier et al. 2011, Figure 1). The DAF are found between 1,900 and 3,400 m of elevation in the north, northwest, and central dry highlands; in the southeast, they occur between 1,500 and 2,200 m (Friis et al. 2010). Various researchers have prepared ecological descriptions of the Ethiopian DAF (Friis and Tadesse 1990, Friis 1992, Woldu 1999, Demissew et al. 2004, Friis et al. 2010). In this article, we have used the description by Friis et al. (2010) because it is the most recent and comprehensive (Table 1 and Figure 2).
Description of the dry evergreen Afromontane forests subtypes in Ethiopia. This table was prepared by extracting information in Friis et al. (2010).
According to several vegetation surveys (Ayalew et al. 2006, Woldemichael et al. 2010, Alemu 2011, Benti 2011, Soromessa and Kelbessa 2014, Woldemariam et al. 2016, Woldearegay et al. 2018, Mohammed et al. 2019, Reshad et al. 2019, Yirga et al. 2019), the following tree species are most characteristic of the Ethiopian DAF: Bersama abyssinica Fresen. (Melianthaceae), Ekebergia capensis Sparrm. (Meliaceae), Juniperus procera Hochst. ex Endl. (Cupressaceae), Olea europaea L. subsp. cuspidata (Wall. and G.Don) Cif. (Oleaceae), and Podocarpus falcatus (Thunb.) R.Br. ex. Mirb. (Podocarpaceae). Therefore, these tree species should be included in DAF restoration programs. Moreover, the potential value of other characteristic tree and shrub species listed in Table 1 should also be considered.
There are about 65 DAF remnant patches in Ethiopia as documented by the Ethiopian Biodiversity Institute and other researchers (Table 2). Of these, 24 are designated as a National Forest Priority Area (NFPA). Based on the forests designated as NFPAs (i.e., protected DAF), the conservation status of the Ethiopian DAF has been determined using indices of “protection” and “connectivity” (Joint Research Centre of the European Commission 2021). Protection was computed as the percentage of the area of the protected DAF to the total area of the DAF-G. Connectivity was computed as a factor of 1) the areal extent of protected DAF, 2) ease of dispersal among the protected forests (i.e., the likelihood of the presence of two protected DAFs within 10 km), and 3) network availability (i.e., protected forest in the nearby ecoregion that could serve as a stepping stone) (Saura and de la Fuente 2017, Saura et al. 2017). The DAF protection and connectivity levels were estimated to be 7.43% and 2.79% respectively, much lower than the national averages for Ethiopia’s ecoregions, which are estimated to be 16.03% and 7.8% respectively (Joint Research Centre of the European Commission 2021). However, these figures were computed based on the area of the DAF-G, which includes areas suitable for DAF as well as for grassland. Hence, the actual levels of DAF protection and connectivity could be a bit higher, but still, much lower than the 17% connectivity proposed for effective conservation of important terrestrial areas (United Nations CBD Secretariat 2019).
The dry evergreen Afromontane forests of Ethiopia.
According to Hodgson et al. (2011), how well forests are managed (i.e., their ecological “quality”) is an even more important indicator of their conservation status. In Ethiopia, DAF occurs coincident with the densest human population. As a result, degradation and deforestation have been ongoing for centuries (Woldu 1999, Bishaw 2001). Furthermore, for the past few decades, DAF management programs (particularly for the protected DAF tracts), have achieved limited success (Mengistu 2003) as corroborated by studies of forest cover change (Table 3). These data (i.e., protection, connectivity, and forest cover change) clearly indicate that the conservation status of the Ethiopian DAF is not good.
Forest cover change reported for some of the dry evergreen Afromontane forests in Ethiopia.
Forest restoration should in no way be an incentive to destroy remnant forests (Chazdon and Laestadius 2016). Protecting and conserving the remnant forests is rather the initial step in forest restoration (DellaSala et al. 2003). Moreover, the presence of natural forest patches nearby greatly helps to ensure the resilience of restored forests by improving connectivity and providing a continuous source of propagules. In turn, the restored forests increase regional connectivity and enhance natural forests’ resilience (Proft et al. 2018). Therefore, future DAF restoration programs in Ethiopia should be augmented with sustainable DAF conservation programs and vice-versa. This is particularly important in the current context because conservation and management of remnant patches receive little attention compared to the massive national campaigns focused on restoration of degraded dry lands through planting trees, shrubs, and herbs.
The Science and Practice of Tropical Forest Restoration
According to the Society for Ecological Restoration (SER 2004), restoration is the process of assisting the recovery of an ecosystem that has been degraded, damaged, or destroyed. In the case of forests, successful restoration returns structure (i.e., species composition, diversity, and relative abundance), function, and resilience that are comparable to those found in undisturbed natural forests that serve as a reference for comparison (Gann et al. 2019). Reducing disturbance, remediation, reclamation, and rehabilitation are all activities along the restorative continuum—activities allied with forest restoration—while afforestation is not (Gann et al. 2019).
Forest restoration is a site-level activity that is different from forest landscape restoration (Chazdon 2017, César et al. 2020) or re-greening (Reij and Winterbottom 2015), which are landscape-level activities. It is also different from reforestation, which usually requires planting trees or sowing tree seeds (Mansourian 2005). Reforestation qualifies as forest restoration only when the goal is to recover as many of the attributes of the pre-disturbance natural forest as possible (Elliott et al. 2013). Although forest restoration is a site-level activity, for the issues of sustainability and climate resilience, it should be planned at the landscape level and within the context of ecoregional conservation (Morrison et al. 2005, Chazdon 2017; Proft et al. 2018).
In practice, restoration employs knowledge from disciplines including ecology, ecophysiology and functional ecology, phytogeography, genetics, engineering, statistical modeling, and biotechnology (Oliet and Jacobs 2012, Walker et al. 2007, Asmelash et al. 2016, Palmer et al. 2016). However, the theoretical bases of restoration are ecological succession, ecological assembly, and ecological disturbance (White and Jentsch 2004, Hobbs et al. 2007). From the viewpoint of forest ecosystem development, ecological succession can be viewed as ecological assembly in progress (Hobbs and Norton 2004, Walker et al. 2007, Bhaskar et al. 2014, Chang and HilleRisLambers 2016).
Succession takes place when species colonize a site (either passively or through intentional introduction) and then perform differently once established (Pickett et al. 1987). Hence, restorationists can control the colonization of a site and also manipulate the performance of the colonizing species to accelerate succession (Luken 1990, Hobbs and Norton 1996, Parker 1997, Bongers et al. 2006). Similarly, restorationists can remove dispersal filters by introducing appropriate species and can modify abiotic filters (e.g., light, nutrients, and water) and biotic filters (e.g., herbivores, parasites, pathogens, pollinators, symbionts) to accelerate community assembly (Hulvey and Aigner 2014, Kraft and Ackerly 2014, Figure 3). Reducing or eliminating disturbances that caused degradation is an important initial step. Replacing these original disturbances with designed disturbances is crucial for driving succession and assembly (Hobbs and Norton 1996, Bongers et al. 2006, Figure 3).
The “priority effect” (i.e., the influence of species that first colonize a site) is also an important consideration in forest restoration (Cortines and Valcarcel 2009, Török et al. 2021). Likewise, facilitation by nurse plants is a concept with a practical application in forest restoration (Bertness and Callaway 1994, Gómez-Aparicio 2009, Gómez-Ruiz et al. 2013, de la Luz Avendaño-Yáñez et al. 2014, Fagundes et al. 2023) including DAF (Aerts et al. 2006 and 2007, Negash and Kagnew 2013, Abiyu et al. 2017).
Based on ecological concepts embedded in and aligned with ecological succession and assembly theories, based on landscape ecology, and documented successful restoration practices, five methods of tropical forest restoration have been recommended. These methods have become widely accepted and are being used as standard methods by multinational organizations that work on forest restoration. Moreover, Chazdon et al. (2021) have grouped a continuum of forest restoration methods into four categories based on the level of intervention each method requires (Table 4, Figure 4).
Description of the five tropical forest ecosystems restoration methods proposed by Elliott et al. (2013).
Attributes of DAF Restoration
According to several studies, dry Ethiopian highland soils rarely harbor seeds of trees characteristic of DAF (Lemenih and Teketay 2004, Teketay 2005a and b, Reubens et al. 2007, Sileshi and Abraha 2014). Hence, DAF restoration from the soil seed bank is almost impossible. The probability that such tree seeds will naturally arrive at open restoration sites is also very low (Lemenih and Teketay 2004). If the seeds are large and animal dispersed, they may not even move beyond forest edges (Cole et al. 2011). For those few seeds that are dispersed, establishment is also a major problem.
Because the DAF occupies areas with a short growing season, strong wind, depauperate soils, low soil organic carbon, considerable cloud cover, and comparatively low levels of solar radiation, there is a high rate of seeding failure and growth of seedlings that can establish is slow. Revegetation is slow compared to that in the surrounding lowland forests (Guariguata 2005, Dalling et al. 2016).
Moreover, as is the case with many tropical landscapes, the DAF could undergo retrogressive succession. This is a long-term process in which forest structure and function (e.g., moisture, carbon, and nutrient cycling) decline due mainly to the effects of nutrient depletion or unavailability (Walker and Reddell 2007, Peltzer et al. 2010). When such forests are cleared, used for cropland, and then abandoned, soil fertility is further reduced, leading to even higher seedling recruitment failure during restoration (Lemenih et al. 2005, Delelegn et al. 2017, Birhane et al. 2018, Asmelash et al. 2021c).
For these reasons, DAF restoration cannot be achieved through natural regeneration alone and trees must be planted. Fortunately, the same mix of trees can be used in most DAF restoration sites because the Ethiopian DAF do not exhibit clear elevational zonation in species composition (Friis 1992).
Compared to other major forest ecosystems in Ethiopia, DAF restoration could significantly be affected by global climate change (Van Breugel et al. 2016). Climate-induced tree dieback has already been recorded in one of the DAF remnants in northern Ethiopia (Mokria et al. 2015). Due to climate change, 20–85% of the DAF-G in Ethiopia could shift to Combretum-Terminalia woodland (CTW) by 2070 (Van Breugel et al. 2016). CTW is an ecoregion dominated by tree species of the genera Combretum and Terminalia (Combretaceae), which are small- to moderate-sized trees with fairly large deciduous leaves (Friis et al. 2010). In the worst-case scenario, only 15% of the land area currently occupied by DAF would continue to be suitable for DAF restoration. Under the best conditions, however, up to 80% could remain suitable. Regardless, there are three possible options for DAF restoration: 1) restoration based on existing reference sites without regard to the effects of climate change, 2) restoration based on anticipated climatic conditions (e.g., CTW), or 3) restoration of a climate-resilient DAF. Of the latter two, restoration advocates would recommend restoring climate-resilient DAF (Corlett 2016), including restoring the largest possible DAF patches at sites that increase connectivity (Proft et al. 2018). Incorporating species from a variety of provenances would be appropriate for restoring resilient forests (Hubert and Cottrell 2007, Bosselmann et al. 2008, Breed et al. 2013, Gann et al. 2019). Establishing a DAF consortium in Ethiopia would be crucial to facilitate the exchange of tree seedlings and seeds for composite provenance planting in a climate-resilient DAF restoration program.
The DAF are found in the DAF-G ecoregion where grasslands are an integral component of the regional plant community. As a result, there may be cases in which DAF restoration would be implemented on natural grasslands or historical grasslands. The Ethiopian dry highland grasslands have been identified to be among the world’s ecosystems at greatest risk of loss due to afforestation (Veldman et al. 2015, Bond et al. 2019). Therefore, DAF restoration needs to be considered from a regional, ecosystem-based perspective to assess the possible impact of forest restoration on native grasslands.
Potential Mechanisms to Improve Tree Seedling Establishment
As has been described above, the restoration of Ethiopian DAF is hardly achievable through natural regeneration alone; planting trees is essential. Therefore, successful seedling establishment defines the success of restoration projects. In this section, we present potential mechanisms to improve tree establishment, survival, and growth in the dry highlands of Ethiopia. We selected these mechanisms for review because their efficacy has been evaluated previously.
Planting Site Preparation and After-planting Care
The initial step in forest restoration on tropical landscapes with multiple resource limitations is to restore the hydrological properties of the site (Walker and Reddell 2007). The construction of soil and water conservation structures (SWCS) is crucial to improve catchment hydrology (Nyssen et al. 2010, Meaza et al. 2022), soil moisture, and nutrient status in the dry highlands in Ethiopia (Demelash and Stahr 2010, Challa et al. 2016, Tolessa et al. 2021). However, little is known if such improvements are sufficient to improve seedling establishment, survival, and growth. Whereas significant positive effects of micro-basin construction have been reported for the survival and growth of some exotic tree species (Derib et al. 2009, Alem et al. 2020), no significant effect was found in the cases of the native species J. procera and O. europaea subsp. cuspidata (Siyum et al. 2019). Therefore, more research is needed to understand the effects of SWCS (including design alternatives) on DAF tree seedlings.
Based on more than two decades’ research on the biology and restoration of DAF tree species, Negash (2021) concluded there are no shortcuts for DAF restoration. Hence, before tree seedlings are planted, planting pits need to be prepared well and watering ponds should be dug where there is no water source near a restoration site. Care of seedlings after planting for at least two years (i.e., manure application, fencing when appropriate, hoeing, weeding, watering during the dry season, and mulching) is crucial (Negash 2021). With such careful preparation and stewardship, up to 100% survival for some of the most important species’ seedlings has been achieved (Table 5), which otherwise exhibit very low rates of survival and growth at natural and experimental field conditions (Aerts et al. 2006, Abebe et al. 2011, Teketay 2011, Asmelash et al. 2019). Hence, site preparation including SWCS construction and seedling after-care should be the standard for DAF restoration. Other mechanisms, discussed below, could substitute for site preparation and after-care when they are found to produce comparable results or are economically more feasible. In fact, several mechanisms may work synergistically to achieve the best results.
Field survival rate of seedlings of some tree species characteristic of dry evergreen Afromontane forests seven months after planting on a degraded dry highland site in central Ethiopia. Survival rates were recorded for quality seedlings raised with desirable traits. Survival rates in the left column were reported by Belude (2007); those in the right column were reported by Tafesse (2007).
Nursery Practices to Improve Successful Restoration
Tree seedlings to be out-planted in the dry highlands in Ethiopia should be raised in a manner that will prepare them to overcome the moisture and nutrient limitations they will face in the field. Generally, growing seedlings in pots rather than as bare root stock increases survival (South et al. 2005, Negash 2021). Raising seedlings with a comparably deeper root (Lobet et al. 2014, Trona et al. 2015), bigger collar diameter (Haase 2007, Grossnickle 2012), and higher root-to-shoot or root-to-dry biomass ratios (Comas et al. 2013) is important to increase field survival. Raising seedlings such that they accumulate more non-structural carbohydrates (NSCs) is also helpful (O’Brien et al. 2014).
The identification and implementation of practices that maximize these desirable traits is crucial. NSC accumulation can be enhanced by increasing the leaf area, photosynthetic rate, and root system size by supplying the seedlings with appropriate levels of nutrients and moisture (Villar-Salvador et al. 2015). Using larger diameter pots has been shown to significantly increase seedlings’ root collar diameter (Grossnickle 2012, Abera et al. 2018). Increasing pot depth was found to significantly increase the root collar diameter and the root-to-shoot and root-to-dry biomass ratios (Gülcü et al. 2010). The practice of root pruning commonly used in nurseries to increase root system size should be avoided as much as possible, because it has been shown to significantly reduce field survival (Negash 2021). The nature of the potting soil and the length of time seedlings are held in the nursery are other factors to consider (Table 6). Further evaluation of the effects and economic feasibility of several nursery practices is warranted.
Important nursery practices for enhancing field survival of some tree species characteristic of the important dry evergreen Afromontane forests. Table prepared by extracting information found in Negash (2021).
Use of Nurse Trees and Shrubs
Competition and facilitation are the two common interactions in plant communities (Young et al. 2001). Facilitation is an interaction resulting in positive effects on at least one of the interacting species (Bertness and Callaway 1994, Brooker et al. 2008). Hence, the use of facilitative techniques in degraded terrestrial ecosystem restoration is gaining prominence (Gómez-Aparicio 2009, Gómez-Ruiz et al. 2013, de la Luz Avendaño-Yáñez et al. 2014).
Although the theory of facilitation has been informing agroforestry practices for a long time, its application in forest restoration is a recent phenomenon (McIntire and Fajardo 2014). Currently, planting with complementary “nurse” species is among five methods of tropical forest restoration (Elliott et al. 2013). In Ethiopia, nurse trees have been used by farmers for centuries in coffee agroforestry. However, there has not been any documented forest restoration project that has incorporated the nurse plantation method. The few studies carried out to evaluate the potential value of using nurse trees or shrubs in Ethiopian DAF restoration have indicated that the use of nurse vegetation could significantly increase survival and growth (Table 7).
Nurse tree/shrub effects reported in the dry highlands in Ethiopia.
Vachellia abyssinica ([Hochst. ex. Benth.] Kyal. & Boatwr.) is among the important nurse tree species for use in DAF restoration. The species’ desirable traits include possession of spines at a young seedling age sufficient to deter herbivores, nitrogen fixation and soil fertility improvement, and drought tolerance (Negash 2021). The survival rate of V. abyssinica on dry highlands in Ethiopia is significantly higher than that of some exotic tree species (Bekele et al. 2021). Another potential nurse plant could be the exotic leguminous tree Acacia decurrens Willd. (Beshir et al. 2022). This species has also been found to produce high-quality charcoal in less than six years (Ferede et al. 2019). Hence, its application as a nurse tree could also significantly increase the economic feasibility of DAF restoration projects. However, serious pest and disease problems associated with using such a nurse plant should not be overlooked (Afework et al. 2024). There are also several other dry highland native Vachellia (formerly Acacia) trees and shrubs with a potential value as nurse species. Large-scale application potential of the nurse trees and shrubs previously identified is also important (Table 7). Croton macrostachyus could be a priority in this regard because it is reported to significantly increase soil fertility, including plant-available phosphorus (Negash 2021). One logical DAF restoration method could be: 1) protect a degraded restoration sites for a few years, 2) remove the re-spouting weedy shrubs, grasses and forbs while leaving trees and shrubs with demonstrated nursing potential, and 3) plant mid- and late-successional tree species’ seedlings.
Arbuscular Mycorrhizal Fungal Inoculation
Arbuscular mycorrhizal fungi (AMF) inoculation has been shown to significantly increase trees seedlings’ field survival (Pouyu-Rojas and Siqueira 2000, Dag et al. 2009, Kapulnik et al. 2010, Karthikeyan and Krishnakumar 2012, Pereira et al. 2021) and growth (Urgiles et al. 2009, Urgiles et al. 2014, Schüßler et al. 2016, Asmelash et al. 2021b). However, inoculation effects depend on the host plant’s responsiveness, the AMF status of the planting material, and the AMF status of the planting sites (Verbruggen et al. 2013). Accordingly, inoculation effects could range from positive to neutral to negative (Johnson et al. 1997).
In the past, few studies have been done to determine the potential benefit of AMF inoculation for DAF restoration in Ethiopia. The simple technique of AMF inoculation resulted in very significant growth improvements of five-month-old seedlings of Cordia africana Lam., an early successional riverine DAF tree species, both on sterile and non-sterile degraded soil. However, seedlings of the mid- and late-successional DAF tree species J. procera and P. falcatus were shown to have low- to negative-AMF responsiveness (Asmelash et al. 2021b). A general trend of a high, medium, and low arbuscular mycorrhizal responsiveness has also been reported for, respectively, early-, mid-, and late-successional tropical tree species elsewhere (Kiers et al. 2000, Zangaro et al. 2003, 2007).
A nursery survey carried out to evaluate the AMF status of planting materials grown for DAF restoration revealed that the nursery stocks were sufficiently colonized (Asmelash et al. 2021a). Regarding the AMF status of planting sites, studies reported contrasting results. While Birhane et al. (2018) and Asmelash et al. (2021c) recorded significant increases in AMF spore abundance and significant reductions in root AMF colonization and infectivity due to site deforestation and degradation, Delegegn et al. (2017) and Birhane et al. (2020) recorded significant reductions of AMF spore abundance and root AMF colonization. Based on these findings, we speculate the AMF inoculation demand of seedlings and sites is low for specific tree species. However, AMF inoculation could be appropriate for selected and AMF-responsive tree species. Therefore, long-term research to investigate the AMF responsiveness of several DAF tree species is crucial. For those AMF-responsive tree species, identifying the most beneficial AMF species or strain(s) is also important.
Biochar Application
Biochar is charcoal used as a soil amendment (McLaughlin et al. 2009, Brockamp and Weye 2020). Biochar application has been shown to significantly increase plant growth and yield (Biederman and Harpole 2013, Kammann et al. 2016, Jeffery et al. 2017), often with a single application (Kätterer et al. 2019, Frimpong et al. 2020). As a result, biochar application could be a promising technology in forest restoration (Thomas and Gale 2015, Palviainen et al. 2020, Grau-Andrés 2021). Considering tropical forests in particular, Román-Dañobeytia et al. (2021), in what is the largest experiment to date, found that biochar application alone and with N-P-K fertilization significantly increased the field survival and growth of tree species with low- and medium-density wood. Biochar’s effects, however, are highly variable and its beneficial effects depend on several factors including application rate and method, feedstock type and pyrolysis temperature/condition, soil conditions, and the plant taxa for which biochar is applied (Thomas and Gale 2015, Joseph et al. 2021). Generally, biochar application is recommended for soils with low cation exchange capacity, soil organic carbon and pH, and with heavier textures (Crane-Droesch et al. 2013)—typical soil conditions in Ethiopia’s dry highlands.
Despite its great potential, there have only been two biochar application studies relevant to DAF restoration in Ethiopia. One of these studies reported no significant application effect on the growth and field survival of O. europaea subsp. cuspidata and Dodonaea angustifolia seedlings (Kayama et al. 2019). On the other hand, the second study reported significant positive effects on the field survival and growth of Faidherbia albida seedlings. Biochar also seemed to enhance the establishment of Vachellia etbica seedlings, but definitive results for this species were compromised by herbivore damage (Kayama et al. 2021). In these two experiments, biochar was applied alone, which may account for observed nonsignificant or low effects. Biochar generally has low N content and its application could require enrichment with organic or inorganic N fertilizers to offset initial N sorption by biochar (Thomas and Gale 2015, Bonanomi et al. 2017).
Conclusion
In this review, we have summarized the scientific and technical issues relevant to DAF restoration. We have also reviewed the literature and gathered data to evaluate the conservation status of the remnant DAF in Ethiopia. We have shown that the conservation status of DAF is not good. Most of these forests have experienced significant reductions in area. Moreover, DAF restoration generally is not possible through natural regeneration alone. Although protection and assisted natural regeneration (ANR) could be very important for DAF restoration, neither is effective unless augmented by tree planting. Framework planting, maximum biodiversity planting, and/or nurse plantation methods are needed, preferably in combination with protection and/or ANR. In some cases, ANR complemented by selective tree species planting could also be a feasible method. In this review, we have presented and discussed five mechanisms for increasing tree seedlings’ establishment, survival, and growth. Proper site preparation and after-planting care is the most effective. The other mechanisms (i.e., modifying nursery practices, incorporating nurse tree and shrubs, inoculating with arbuscular mycorrhizal fungi, and applying biochar) also have potential applications in DAF restoration. However, they cannot replace proper planting site preparation and after-care stewardship. The application of the five mechanisms complementarily could be most effective. However, there are still more questions than answers regarding the effectiveness of these five mechanisms. We hope this review will motivate researchers to engage further to answer some of the important questions.
Socioeconomic factors such as reducing deforestation, avoiding overgrazing, and using exclosures could also help DAF restoration. Successful restoration will require comprehensive and integrated approaches incorporating both ecological and social interventions.
Author Contributions
This project was conceptualized primarily by FA, with contributions by MMR. FA completed the literature review and wrote the original draft; MMR reviewed the draft.

Footnotes
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